Gammaridean Amphipoda
of Australia, Part III:
The Phoxocephalidae
J. LAURENS BARNARD
and
MARGARET M. DRUMMOND
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 245
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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 245
Gammaridean Amphipoda
of Australia, Part III:
The Phoxocephalidae
J. Laurens Barnard
and Margaret M. Drummond
SMITHSONIAN INSTITUTION PRESS
City of Washington
1978
A B S T R A C T
Barnard, J. Laurens, and Margaret M. Drummond. Gammaridean Amphipoda
of Australia, Part III: The Phoxocephalidae. Smithsonian Contributions to
Zoology, number 245, 551 pages, 269 figures, 1978.?Phoxocephaiidae are found
to be the most diverse family of benthic marine amphipods in southern Australia.
This study reports on 88 species in 26 genera. Most of the taxa are described as
new. The Phoxocephalidae of the world are reorganized by the description of 8
new subfamilies and the removal of all but 2 species from Paraphoxus either to
new genera or by indication that new genera must be described. Some taxa of
Australian phoxocephalids are shown to be the most primitive known to exist,
especially Pontharptnia and a new subfamily Tipimeginae. Lines of evolutionary
deployment radiate outward from Australia to South America and then northward
into the northern hemisphere. The expected close relationship to urothoids and
pontoporeids is briefly discussed; in addition, an affinity to Ponto-Caspian gam-
maroids is also explored. Extraordinarily dense populations of phoxocephalids are
found in embayments of Australia, especially Western Port and Port Phillip Bay.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded
in the Institution's annual report, Smithsonian Year. SERIES COVER DESICN: The coral Montastrea
cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data (Revised)
Barnard, Jerry Laurens, 1928-
Gammaridean Amphipoda of Austrialia.
(Smithsonian contributions to zoology, no. 103, 139, 245)
Bibliography: pt. 1, p. 318; p i 2, p. 135; pt- 3, p.
Includes index.
Contents: Pt. 3.
Barnard, J. L. and Drummond, M. The Phoxocephalidae.
1. Amphipoda?Australia. 2. Crustacea?Australia. I. Drummond, Margaret M. II. Title.
III. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 103, etc.
QL1.S45 no. 103. etc. 591'.08s [595'.371] 72-601600
Contents
Page
Introduction 1
Acknowledgments 1
Organization of the Study 6
Materials 6
Methods 7
Mechanics of Presentation 7
Terms 8
Text Abbreviations 8
Figure Abbreviations 8
Counts and Measurements 9
Nomenclature 11
Presentation of Species Data 11
Illustration Technique 12
Taxonomic Characters of Phoxocephalidae 15
Evaluation of Females 16
Standard Description of Male Phoxocephalid 26
Male Phoxocephalids 26
Minor Attributes 27
Senility 29
Summary of Taxonomic Evaluations 29
Phoxocephalid Environment 29
Phoxocephalidae in Australia 29
Biogeography of Phoxocephalidae 31
Phoxocephalid Evolution 34
Origin of Phoxocephalidae 34
Evolutionary Descent in Phoxocephalidae 37
PHOXOCEPHALIDAE 39
Key to the Subfamilies of Phoxocephalidae 39
PONTHARPINIINAE 40
Pontharpinia Stebbing 40
Pontharpinia pinguis (Haswell) 41
TIPIMECINAE 46
Key to the Genera of Tipimeginae 46
Tipimegus, new genus 47
Key to the Species of Tipimegus (Adults) 49
Tipimegus thalerus, new species 49
Tipimegus dinjerrus, new species 55
Tipimegus kangulun, new species 58
Tipimegus kalkro, new species 64
Tipimegus stebbingi (J. L. Barnard) new combination 70
Tipimegus species 4N 70
Tipimegus species 5N 70
iii
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
IV
Page
Booranus, new genus ' '
Key to the Species of Booranus '2
Booranus weemus, new species '2
Booranus tikeri, new species 76
Booranus wangoorus, new species 81
Trichophoxus K. H. Barnard 85
Trichophoxus capillatus K. H. Barnard 86
Waitangi Fincham 86
Waitangi rakiura (Cooper and Fincham) 87
BROLGINAE 87
The Brolgin Group of Genera 87
Relationships of the Brolgin Group to Metaphoxus and Phoxoce-
phalus 89
Key to the Genera of Brolginae 89
Mandibulophoxus J. L. Barnard 90
Key to the Species of Mandibulophoxus 91
Mandibulophoxus uncirostratus (Giles) 91
Mandibulophoxus gilesi J. L. Barnard 91
Cunmurra, new genus 92
Cuntnurra itickerus, new species 93
Brolgus, new genus 96
Keys to the Species o/ Brolgus 97
Brolgus tattersalli (J. L. Barnard), new combination 98
Brolgus millinus, new species '04
Brolgus mahmak, new species '08
Brolgus tavelus, new species " 3
Brolgus koongarrus, new species "**
Elpeddo, new genus " 8
Elpeddo kaikai, new species " 9
Ganba, new genus '24
Ganba pellati, new species '25
Kuritus, new genus '29
Kuritus nacoomus, new species '30
Wildus, new genus '33
Key to the Species of Wildus '34
Wildus thambaroo, new species '34
Wildus mullokus, new species '38
Wildus} fuegiensis (Schellenberg), new combination '44
Wildus? waipiro (J. L. Barnard), new combination '44
Paraphoxus Sars 144
Paraphoxus oculatus (Sars) 145
Paraphoxus} simplex (Gurjanova) 145
Allocation of Species in Paraphoxus 145
LEONGATHINAE 146
Leongathus, new genus 146
Leongathus nootoo, new species 147
JOUBINELLINAE 152
Key to the Genera of Joubinellinae 152
NUMBER 245
Page
Joubinella Chevreux 153
Key to the Species of Joubinella 153
Joubinella species 1 154
Joubinella species 2 154
Matong, new genus 154
Matong matong, new species 155
Kotla, new genus 161
Kotla batturi, new species 161
Yammacoona, new genus 166
Yammacoona kunarella, new species 167
PARHARPIMINAE 174
Parharpinia Stebbing 175
Key to the Species of Parharpinia 176
Parharpinia villosa (Haswell) 176
Parharpinia warte, new species 185
Protophoxus K. H. Barnard 189
Protophoxus australis K. H. Barnard 190
BIRUBIINAE 190
Key to the Genera of Birubiinae 191
Birubius Barnard and Drummond 191
The Evolutionary System in Birubius 193
Key to the Species of Birubius (Females) 193
Aids to Identification in Birubius 199
1. Birubius panamunus Barnard and Drummond 201
2. Birubius lorus, new species 207
3. Birubius nammuldus, new species 212
4. Birubius myallus, new species 216
5. Birubius kareus, new species 222
6. Birubius apart, new species 228
7. Birubius cartoo, new species 234
8. Birubius thalmus, new species 239
9. Birubius muldarpus, new species 242
10. Birubius gallangus, new species 247
11. Birubius mayamayi, new species 251
12. Birubius wirakus, new species 260
13. Birubius chintoo, new species 264
14. Birubius karobrani, new species 268
15. Birubius booleus, new species 272
16. Birubius babaneekus, new species 276
17. Birubius gelarus, new species 281
18. Birubius quearus, new species 285
19. Birubius narus, new species 288
20. Birubius gambodeni, new species 292
21. Birubius maamus, new species 299
22. Birubius lowannus, new species 304
23. Birubius kyeemus, new species 309
24. Birubius batei? (Haswell), new combination 312
25. Birubius kokorus, new species 320
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
VI
Page
26. Birubius kinkxis, new species 32J
27. Birubius munggai, new species 329
28. Birubius ularitus, new species 334
29. Birubius eleebanus, new species 338
30. Birubius jirrandus, new species 313
31. Birubius yorlunus, new species 351
32. Birubiu eake, new species 357
33. Birubius kabbulinus, new species 362
34. Birubius taldeus, new species 365
35. Birubius yandus, new species 370
36. Birubius maldus, new species 375
37. Birubius wulgaru, new species 380
Yan, new genus 388
Key to the Species of Yan 390
Yan tiendi, new species 390
Yan errichus, new species 393
Tickalerus, new genus 396
Tickalerus birubi, new species 398
Kulgaphoxus, new genus 402
Kulgaphoxus borralus, new species 404
Kulgaphoxus cadgeeus, new species 408
Microphoxus J. L. Barnard 414
Microphoxus minimus J. L. Barnard 415
Metharpinia Schellenberg 415
Metharpinia longirostris Schellenberg 416
PHOXOCEPHALINAE 416
Key to the Genera of Phoxocephalinae 416
Phoxocephalus Stebbing 417
Key to the World Species of Phoxocephalus 418
Phoxocephalus bassi (Stebbing) 418
Phoxocephalus kukathus, new species 422
Phoxocephalus tunggeus, new species 427
Phoxocephalus rupullus, new species 431
Phoxocephalus burleus, new species 435
Phoxocephalus keppeli, new species 441
Phoxocephalus species 441
Phoxocephalus holbolli (Kr0yer) 441
Phoxocephalus homilis J. L. Barnard 441
Phoxocephalus kergueleni (Stebbing) 441
Phoxocephalus regium K. H. Barnard 442
Phoxocephalus tenuipes Stephensen 442
Jerildaria, new genus 442
Jerildaria joubiphoxus, new species 443
Leptophoxoides J. L. Barnard 446
Leptophoxoides molaris J. L. Barnard 447
Leptophoxus Sars 447
Leptophoxus falcatus falcatus (G. O. Sars) 448
Leptophoxus falcatus icelus J. L. Barnard 448
NUMBER 245 yjj
Page
Metaphoxus Bonnier 448
Key to the Species of Metaphoxus 449
Metaphoxus tuckatuck, new species 449
Metaphoxus yaranellus, new species 456
Metaphoxus mintus, new species 460
Metaphoxus frequens J. L. Barnard 465
Metaphoxus fultoni (Scott) 465
Metaphoxus pectinatus (Walker) 465
Metaphoxus tulearensis, new species 466
Metaphoxoides Ledoyer 466
Key to the Species of Metaphoxoides 467
Metaphoxoides picardi Ledoyer 467
Metaphoxoides zavorus, new species 467
Diogodias, new genus 467
Key to the Species of Diogodias 468
Diogodias littoralis (Cooper and Fincham), new combination 468
Diogodias longicarpus (Ledoyer), new combination 468
Diogodias platyrostris (Ledoyer), new combination 468
Vasco, new genus 468
Vasco brevidactylus (Ledoyer), new combination 469
Hopiphoxus, new genus 469
Hopiphoxus simillimus (J. L. Barnard), new combination 470
Rikkarus, new genus 470
Rikkarus lea, new species 471
Japara, new genus 474
Japara papporus, new species 475
Kondoleus, new genus 480
Kondoleus tekin, new species 481
Limnoporeia Fearn-Wannan 487
Key to the Species of Limnoporeia and Uldanamia 488
Limnoporeia kingi Fearn-Wannan 489
Limnoporeia maranowe, new species 489
Limnoporeia yarrague, new species 495
Limnoporeia woorake, new species 501
Limnoporeia ungamale, new species 506
Limnoporeia wakkine, new species 511
Limnoporeia kalduke, new species 517
Uldanamia, new genus 522
Uldanamia pillare, new species 522
HARPINIINAE 528
The Harpiniin Group of Genera 529
Key to the Genera of Harpiniinae 529
Coxophoxus J. L. Barnard ? ? 530
Coxophoxus hidalgo J. L. Barnard 530
Coxophoxus coxalis (K. H. Barnard) 530
Basuto, new genus 530
Basuto stimpsoni (Stebbing), new combination 531
Proharpinia Schellenberg 531
vJjj SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Page
Heterophoxus Shoemaker 532
Pseudharpinia Schellenberg 533
Pseudharpinia dentata Schellenberg 534
Harpiniopsis Stephensen 534
Harpinia Boeck 535
Appendix: Australian Samples and Localities 537
Literature Cited 539
Index 545
Gammaridean Amphipoda
of Australia, Part III:
The Phoxocephalidae
J. Laurens Barnard
and Margaret M. Drummond
Introduction
The Phoxotephalidae in Austrialian marine shal-
lows constitute the largest familial element of Am-
phifxxla discovered so far on that continent. Most
of the taxa are newly described. This study reports
on 88 species in 26 genera, comprising about 40
percent of the known taxa worldwide. Both Aus-
tralia and the world in general remain poorly ex-
plored for this group.
Australia appears to have the most primitive
known assemblage of phoxocephalids as represented
by: the endemic Pontharpiniinae; its satellite the
Tipimeginae, composed of several genera from both
Australia and New Zealand; the Brolginae, com-
posed of five genera with more advanced members
being found in South America and the northern
hemisphere; and the Birubiinae and Parharpini-
inae, with lines of dispersal towards the American
continents.
Almost all of the phoxocephalid characters or
generic attributes have been found, at least in rudi-
mentary form, in the Australian fauna. The only
other major center of distinctive attributes in the
J. Laurens Barnard, Department of Invertebrate Zoology,
National Museum of Natural History, Smithsonian Institu-
tion, Washington, D.C. 20560. Margaret M. Drummond,
Marine Pollution Studies Group, Fisheries and Wildlife Divi-
sion, Ministry for Conservation, 605 Flinders Street Extension,
Melbourne, Victoria 3000 Australia.
family is found in the subantarctic islands or the
cold-temperate part of South America where the
Proharpinia-Pseudharpinia-Heterophoxus (Harpini-
inae) center of distribution is located. Phoxocepha-
lids in other parts of the world, though known only
sketchily, appear to be advancements from or de-
scendents out of these two centers, Australia and
Subantarctica. Although these two foci may have
been primary centers of evolution in the group,
they might simply be relict areas in which certain
primitive taxa have been preserved.
The Phoxocephalidae appear to have their closest
morphological counterparts not only in the ponto-
poreid-urothoid-haustoriid family groups but also
in the primitive pontogammarid groups of the
Ponto-Caspian basin. The latter relationship, how-
ever, is strongly disjunct both morphologically and
geographically in the present time. There may have
been a closer relationship geographically in times
of Pangaea.
The immense diversity of amphipodan species in
the bays of Victoria, especially that of Western
Port (= Westernport), deserves special mention in
the context of this study.
ACKNOWLEDGMENTS.?In Australia our study has
been helped in many ways by a number of individ-
uals and organizations. In particular, we acknowl-
edge the debt to Mr. A. Dunbavin Butcher, Deputy
Director of Conservation in Victoria, who, as Direc-
tor of Fisheries and Wildlife, was in large part
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pan STEP
BARRCW ISLAND^
Indian
Ocean [
HOUTMAN
ABROLHOS ISLANDS *
GARDCN ISLA
SCALE 1 : 10 001 000
BASS STRAIT
FIGURE 1.?Map of Australia showing major collecting localities.
responsible for initiating the vital surveys in West-
ern Port and Port Phillip Bay. We have appre-
ciated his continued interest in our study. The
Ministry of Conservation of Victoria has listed this
present monograph as Publication Number 6 of
their Environmental Studies Series.
These surveys were supported by subventions
from the Government of Victoria and from indus-
try. The Crib Point Benthic Survey, confined to a
small area around the newly-constructed refinery
and productive of an enormous amphipod collec-
tion, was financed entirely by British Petroleum
Australia Ltd. Four firms, British Petroleum Aus-
tralia Ltd., The Broken Hill Proprietary Co. Ltd.,
Esso Australia, Ltd., and John Lysaght (Australia)
Ltd. (all now represented at Western Port) spon-
sored, jointly with the Victorian Government, the
more extensive Western Port survey of 1973-74.
The Port Phillip Environmental Study was a joint
project of the Melbourne and Metropolitan Board
of Works and of the Victoria Department of Fish-
eries and Wildlife.
Within the Division of Fisheries and Wildlife,
appreciation is due: Dr. Alistair J. Gilmour, As-
sistant Director of Marine Pollution Studies, who
has given us much practical help and advice; Dr.
Gary C. B. Poore of the Marine Pollution Studies
Group, who placed at our disposal Phoxocephalidae
from the Port Phillip Bay Environmental Study
and supplied all station data for that large collec-
tion; Mrs. Danuta Karpow, who, in addition to
assisting ably in the laboratory, has collated the
NUMBER 245
BASS
luttt t ?  laclain Stall*!* 121-127
laid 2 : laclaOs Statical 221-233
laid 1 : laclari** Statiaa* 333-331
FICURE 2.?Map of Port Phillip Bay, Victoria, showing collecting localities.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
LESEMB
BASS STRAIT
FIGURE 3.?Map of Western Port, Victoria, showing collecting localities.
species list; and the skippers and technical crews
of the Division's Research vessels, whose care in the
taking and initial treatment of samples resulted in
the assembling of a first rate collection for us to
study.
We wish to thank Mr. John McNally, Director of
the National Museum of Victoria, Dr. D.J.G. Grif-
fin of the Australian Museum, Dr. R. W. George of
the Western Australia Museum, and Dr. David C.
Lee of the South Australian Museum for providing
facilities for examining their collections, and for
loan of material; Dr. L. Collett of the New South
Wales State Fisheries for loan of material; and Dr.
D.J.G. Griffin, Dr. Roger Lincoln, British Museum
(Natural History), Dr. Torben Wolff, Universitetets
Zoologiske Museum, Copenhagen, and Dr. Charles
Pettit of the Manchester University Museum,
United Kingdom, for help in the search for ma-
terial. We are also indebted to Dr. A. A. Fincham,
British Museum (Natural History), for an advance
copy of his study (1977) on Waitangi.
We are grateful to Miss Helen Fisher of The
Australian Museum and Miss Cathy Drummond of
the Fisheries Branch of the Department of Lands
NUMBER 245
HASTIN6S
CRIB POINT
BENTHIC SURVEY AREA
1115
1114 S?r?.T
-, At ?* kJ ?? ** *?
cm \ . ? ? ? ? ?
X
UM (Mat ^? -~*" \
FICURE 4.?Map of Western Port, Victoria, showing Crib Point Benthic Survey collecting
localities.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
and Forests, New South Wales, for the opportunity
to examine phoxocephalid collections and relevant
data from, respectively, the Shelf Benthic Survey
and the New South Wales State Fisheries Estuarine
Benthic Survey.
The Australian maps were prepared by the Cen-
tral Draughting Section of the Ministry for Con-
servation in Victoria, under the direction of Mr.
D. J. Lloyd. The Draughting Section of the Mel-
bourne and Metropolitan Board of Works kindly
loaned, for copying, the original map of Benthic
Sampling Stations in the Port Phillip Bay Environ-
mental Study.
The Smithsonian Institution fund for support of
the study of taxomony in Indian Ocean biota fi-
nanced the work of Wilma A. Findley, Carolyn L.
Cox, and Biruta Akerbergs, who inked our draw-
ings and readied them for publication. Charline M.
Barnard and Anne B. Cohen prepared the bibliog-
raphy and collated the manuscript. We thank Dr.
E. L. Bousfield, National Museum of Canada, for
offering valuable suggestions and support during
the five years consumed in this study. On behalf of
the Smithsonian Institution the senior author grate-
fully acknowledges the generous gift of duplicate
specimens tendered by the various sources. Our
editor, Barbara T. Spann, contributed many im-
provements to our work.
ORGANIZATION OF THE STUDY.?Because a large
percentage of the world taxa in Phoxocephalidae
are newly described in this treatise, the opportunity
is taken to present a revision of the family by creat-
ing new keys and diagnoses for all genera and by
discussing the important elements of morphological
diversification and the major evolutionary events
that have occurred. This synthesis replaces and re-
vises the phoxocephalid compilation given by J. L.
Barnard (1960). Because few studies on.elements of
the Phoxocephalidae have appeared since 1960, the
replication of keys to species in several non-Austral-
ian genera such as Harpinia, Harpiniopsis, and
Heterophoxus is unnecessary. Keys and diagnoses of
all genera, lists of all known species in heavily re-
vised genera, and updated lists in unmodified
genera are given, however, so that a broad per-
spective to generic level can be presented.
The main contributions to phoxocephalid taxon-
omy since 1960 include the description of Limno-
poreia by Fearn-Wannan (1968), Coxophoxus by J.
L. Barnard (1966a), Metaphoxoides by Ledoyer
(1968), a new species of Metaphoxus by J. L. Bar-
nard (1967) (here relegated to a new genus), a new
species of Paraphoxus by J. L. Barnard (1972b)
(here relegated to a new genus), new species of
Metaphoxus by Ledoyer (1973) (here relegated to
new genera), other new species described by J. L.
Barnard (1962, 1964a, 1972), a review of Mediter-
ranean phoxocephalids by Karaman (1973) and a
review of western Atlantic species by Bousfield
(1973).
MATERIALS.?The bulk of our material comes
from three Victorian surveys, conducted in Western
Port and Port Phillip Bay l>etween 1964 and 1974
by the Department of Fisheries and Wildlife (since
1973 a Division of the Ministry for Conservation in
Victoria). These are: the Crib Point Survey (CPBS),
carried out in Western Port in 1965, preceded by a
small preliminary survey in 1961 and followed up
with seasonal samplings at five selected stations
until 1970; the Port Phillip Bay Environmental
Study (PPBES), 1968-72; and the Western j>ort Bay
Environmental Study (WPBES), 1973-74.
In addition to this material we also had at our
disposal, through the good offices of The Australian
Museum and the New South Wales State Fisheries
respectively, phoxocephalid collections from two
surveys recently undertaken in New South Wales:
Shelf Benthic Survey (SBS), in waters off Sydney;
and New South Wales State Fisheries Estuarine
Benthic Survey (EBS) in the Georges River, Jervis
Bay, Botany Bay, and Lake Macquarie.
We have gleaned all available material from the
Australian Museum, Sydney; the Western Austral-
ian Museum, Perth; the South Australian Museum,
Adelaide; and the National Museum of Victoria,
Melbourne.
Ralph H. Miller (RHM), of the University of
Melbourne, loaned us a large private collection of
plankton samples taken over the period 1967-70 in
the Crib Point area of Western Port. Some of our
personal collections from the Western Australian
intertidal (JLB AUS), and from New South Wales
(MMD), also have contained phoxocephalids in-
cluded in this report.
Through examination of the more than 5300
phoxocephalids in warm-temperate Australia pro-
vided from these sources, we have identified a large
number of species, but this is probably a very small
proportion of those still to be discovered in Aus-
tralian waters. When one considers that 90 percent
NUMBER 245
of our material has come from sampling programs
confined mainly to a few protected embayments,
and that from the one third million square miles
of open-sea coastal shelf around Australia we have
seen fewer than two score samples, it may be antici-
pated that, as different areas are explored, substan-
tial additions will be made to the phoxocephalid
taxa.
A condensed list of samples comprises the Ap-
pendix; station localities in Western Port and Port
Phillip Bay are presented in Figures 2, 3, and 4;
and other localities from which material has come
are shown in Figure I. The position of West Island,
near Kangaroo Island, is shown to correct the er-
roneous arrow in J. L. Barnard (1972a:3, fig. 1).
METHODS.?In the quantitative surveys of West-
ern Port (WPBES) and Port Phillip Bay (PPBES)
most of the l>ottom samples were collected with a
0.1 m2 Smith-Mclntyre benthic grab, operated from
one of the Division's research vessels. Intertidal
flats had to IK- sampled at high tide from a flat-
bottomed barge. Additional dredgings were carried
out at some of the Western Port stations for quali-
tative appraisal.
Immediate gross sorting of each sample was ef-
fected on hoard the vessels by washing the contents
of the grab through a graded series of sieves (the
minimum mesh aperture being 1.0 mm), and pre-
serving the residues in a 5 percent neutral solution
of formaldehyde in seawater for storage. Subsequent
separation of organisms from sediment and the
final sorting of organisms into higher taxa was
done in the laboratory, by hand, with the aid of
magnifying lamp and stereomicroscope. Sorted ma-
terial was stored in 70 percent ethyl alcohol.
Samples collected in Port Phillip Bay (Figure 2)
were obtained on intersections of a simple grid sys-
tem (PPBES). The 52 stations of the Western Port
Survey of 1965 (CPBS) were disposed along a num-
ber of radii centering on Crib Point, and concen-
trated within an area of 915 hectares (Figure 4).
The WPBES (1973-74) samples were obtained from
stations more widely spread, and randomly selected
from within three habitats: intertidal flats, shallow
sublittoral areas to a depth of 5.5 m, and deeper
sublittoral areas (Figure 3). Each sample was ob-
tained from a station at which numerous environ-
mental conditions were also recorded, but these
data and the ecology of the phoxocephalids will be
reported in other media.
Mechanics of Presentation
The format of this study is designed to include
an overview of world Phoxocephalidae to generic
level. A phyletic system is proposed (page 37) and
the text is presented in the order of that system
(with the exception of the genera Rikkarus, Japara,
Kondoleus, and Limnoporeia, which, for ease of
discussion are presented in a different order). Non-
Australian taxa are intermingled with Australian
taxa to facilitate discussion of the interrelationships
of higher taxa. Within diverse genera the Austral-
ian (and New Zealand) species are presented first
in phyletic order, followed by an alphabetical ar-
rangement of exotic species. Presentation of the lat-
ter is limited to their synonymy and distribution.
At the present time the knowledge of minute mor-
phology of the exotic species necessary to incor-
porate them into a world phyletic system is un-
available.
By arranging the species of large genera such as
Birubius (37 species) into clustered phyletic systems,
the discussion of interrelationships has been con-
densed. In each cluster one species is selected as the
model to which the following species are compared
(Figure 82). Later clusters are compared to antece-
dent clusters by groups. Convergence and replica-
tion of evolutionary trends occur in numerous clus-
ters otherwise remotely related to each other. This
occurrence is so common that we have appended an
exposition of unusual replications in Birubius as
an example ("Aids to Identification in Birubius").
This follows the system designed for Paradexamine
by J. L. Barnard (1972b) and might aid future stu-
dents of evolution in this group to detect possible
pathways of descent by early recognition of con-
vergences.
All world genera are rediagnosed and described
so as to match the new diversity of detail necessary
to the elaboration of Australian taxa. Generic
diagnoses and descriptions are produced according
to a model suitable to the Australian group. The
programming lists a set range of alternatives for
each character. If data relative to this character are
not available, the alternatives are listed in brackets.
Keys to all subfamilial groups, genera, and their
species are newly constructed, but new keys are not
provided for groups not studied since 1960.
Several new genera comprising species groups
heretofore assigned to Parnphoxus will be"*described
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
in a later paper.
Each species presentation begins with "Descrip-
tion," based on females where possible, a condensed
description of the male and juvenile when known,
and usually paragraphs with the headings "Obser-
vations" (concerning characters not included in the
description and minor anomalies exhibited by the
specimens described) and "Variations" (noting, in
most cases, normal deviations evidenced by any of
the specimens associated with the taxon). Often a
section under the heading "Illustrations" is in-
cluded to explain anomalies, omissions, and com-
parisons pertaining to the figures. Designations
under the headings "Holotype," "Type-Locality,"
and "Voucher Material" follow. Paragraphs under
"Relationship," (and occasionally "Remarks," hav-
ing reference to problems not concerned with rela-
tionship) and condensed terminal sections under
"Material" and "Distribution" complete the presen-
tation. Further explanation of this format is con-
tained under "Presentation of Species Data."
TERMS.?To save space and cut printing costs,
various grammatical and syntactical devices have
been used. For example, we have often used sub-
stitute terms such as "long" for "elongate," and
terms with "ad" endings such as "laterad" or
"ventrad," refering to the flow or direction of a
character, e.g., setae "extending ventrad," meaning
setae extending in a ventral direction.
The term "ordinary" is used frequently to con-
dense what would otherwise be laborious and often
replicated descriptions of certain characteristics bet-
ter visualized than described. Whenever the term is
used the reader is advised to turn to illustrations to
see the average condition implied and to cross check
diagnoses of other genera to see alternatives. For
example, the alternatives to "article 2 of antenna 1
ordinary" are: "especially shortened" and "espe-
cially elongate." The only alternative to "pereopod
5 ordinary" is "pereopod 5 especially small," and
the only alternative to "pereopod 5 article 3 ordi-
nary" is "article 3 greatly enlarged." The alterna-
tive to "dactyl normal" is "dactyl vestigial." The
terms "normal" and "ordinary" are used in the con-
text of our present knowledge of phoxocephalids;
they may require modification after future discov-
eries.
The word "medium" is used to denote a condi-
tion halfway between extremes; thus a medium
spine is neither "elongate" nor "shortened." The
terms are employed only where alternatives are dis-
tinctive and mostly self-explanatory. The phrase
"continuously spinose" is applied to rami of uro-
pods 1-2 to denote the presence of spines more or
less evenly distributed on the apical third of each
ram us.
The term "Mark" (M.) followed by a number
0-100 refers to a point on an appendage, article, or
ramus, the distance from which point to the base
of the structure is expressed as the jKMtentage of the
total length of that structure. For example, a spine
at Mark 70 (or M. 70) on the outer ramus of uro-
pod 3 lies 0.70 (or 70 percent) of the distance from
base to apex of the stated ramus.
TEXT ABBRF.VIATIONS
AM The Australian Museum, Sy?lncy
CPBS Crib Point Bcuthir Survey. Western Port
EBS New South Walr* State Kishnir* F.Muarine Ben-
thic Survey
JLB AUS collected by J. I.. Barnard
M. Mark, nee definition herein on page 8.
MMI) collected by Margaret M Drummond
NMV National Museum of Victoria, Melbourne
NSW New South Wale*
PPBF.S Port Phillip Bay Environmental Study
RHM samples from the plankton collection of Ralph H.
Miller, University of Melbourne
SAM South Australian Museum, Adelaide
SBS Australian Museum Shelf Bcuthir Survey
T total
WAM Western Australian Museum, Perth
WPBES Westernport Bay Environmental Study
FIGURE ABBREVIATIONS
fused in specimen drawings)
A
B
C
D
E
F
G
H
I
J
K
L
M
N
O
P
0.
R
antenna
prebuccal from lateral
head
dactyl of pereopod
coxa
accessory flagellum
gnathopod
peduncle
inner plate or ramus
right lacinia mobilis
spine, seta, or denticle
lower lip
mandible
molar
outer plate or ramus
pereopod
cuticle
uropod
5 maxilliped
T telson
U upper lip from anterior
V palp
W pleon (pleonites 1-3
bearing epimera;
pleonites 4?6, also
referred to as
urosomites 1-3,
comprising urosome;
often shown with
attached structures:
telson and uropods
1-3)
X maxilla
Y see legend under
illustration
Z calceolus
NUMBER 245
n
o
P
r
s
t
u
V
y
oblique
opposite
in situ
reduced
setae removed
right
part
ventral
enlarged
Lower case letters on the left side of labels denote
specimens cited in the legends and voucher material
in the text; lower case letters on the right side of
labels (or affixed to the drawings as is often the case
with "e" and "s") indicate the following:
a alternative view
c eyes omitted
d dorsal
e broken
/ flat
h half
i medial
; unenlarged
/ lateral
tn abnormal
For other lower case letters see definition in legend.
Where space does not allow a horizontal alinement
of the label, the resulting vertical arrangement has
the same order of elements top to bottom as found
in the horizontal from left to right.
COUNTS AND MEASUREMENTS (Figure 5).?Body
length of amphipods in this study is computed from
a lateral parabolic line drawn from the apex of the
rostrum through the middle of the head, along the
dorsal gut margin to the posterior limit of pleonite
6 at its posteroventral corner. Measurements are
made generally in tenths of millimeters, but in
species where numerous specimens of very similar
sizes are compared, the level is extended to hun-
dredths of millimeters.
Length of head is measured from apex of rostrum
through the midaxis to a posterior point precisely
tangential to the posterior limit of the dorsopos-
terior corner of the head as seen in lateral aspect.
This length is quoted herein as a percentage of the
total body length and includes that part of the head
telescoped into body segment 1. The observation of
total body length is not highly satisfactory because
of segmental telescoping after death. Body size,
length, or other precise measurements are not
highly reliable and are not used in amphipod
taxonomy as extensively as in other fields of study.
Ocular pigmentation is included in the descrip-
tion of each species as a weak but potentially valu-
able characteristic. Most of our materials were pre-
served originally in formaldehyde and then
transferred to alcohol and held one or more years
before examination. Most of the CPBS specimens
were held up to four years in formaldehyde before
being transferred to alcohol. The eyes are generally
reddish brown. PPBES and WPBES specimens, on
the other hand, were held only a few weeks in for-
maldehyde before being transferred to alcohol. The
eye pigment is dark purplish-black in these speci-
mens. Occasional species retain ocular pigment for
many years, others do not. Eyes termed as "stained"
are evenly but translucently pigmented. A standard
cycle of preservation and examination as described
by J. L. Barnard (1970) might reveal good descrip-
tive differences in ocular pigment in the Australian
phoxocephalids.
Spine formulas on articles 4 and 5 of antenna 2
concern the facial spines, quoted from distal to
proximal. On article 4 in certain species, the first
spine group, often restricted to one spine, occurs at
the dorsodistal corner and is often attached medi-
ally; in other species this spine is absent or is joined
laterally by other spines forming the first group.
The ventral counts of setae on article 4 do not in-
clude the basal plusetules which in most phoxo-
cephalids occur medially but which in our illustra-
tions are heavily inked for clarity rather than being
reduced to dots to denote their medial position.
On article 5 the dorsal margin is termed naked
if no armaments other than one dorsodistal setule
group are present. Facial spines near the dorsal
margin are usually easy to distinguish but the api-
calmost facial spine is often shifted so close to a
pair of ventrodistal spines that the three appear to
be in one group. In Birubius, if three of those
ventrodistal spines are present, one is removed to
the facial formula, usually quoted, for example,
1-2-2.
Aesthetascs are usually present on the primary
fiagellum of antenna 1, though not on all articles.
Commonly they are borne on all except proximal
articles 1-3 and the terminal article (Birubius
mayamayi, Brolgus tattersalli, Parharpinia villosa,
etc.). In other species they are confined to the distal
articles, or may be absent altogether; and in one
species they are present on all articles. The descrip-
tions mainly differentiate between gross abundance
or size of aesthetascs, without further elaboration of
detail.
In descriptions of calceoli on antenna 2 of males,
a condensed flagellar formula, e.g., "28, 2, 3, 5, 7 . . .
27," includes the total number of flagellar articles
first, followed by the sequence of segments having
attached calceoli, beginning with article 2 and ob-
serving the progression 2, 3, 5, 7, up through article
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 5.?Location of measurements for body and separate parts (see "Figure Abbreviations" for
key to symbols not defined in this legend); use of term "Mark" (on R3 the point marking end of
article 1 of inner ramus designated M 100; length of outer ramus expressed as a percentage of
the line that measures the length of inner ramus article 1; in this case Af 84); five kinds of molars
(designations A-E correspond to developmental stages described on page 19); setal count on arti-
cle 2 of gnathopods 1-2 and pereopods 1-2 (Y ? article just described, left side of Y = anterior,
L ? long, 5 ?= short, NO = not counted in formulae).
27. When a variation in this formula is added, the
first term (total number of articles) and/or the final
term (last article bearing a calceolus) are omitted if
they are identical to their counterparts in the orig-
inal statement of the formula. Usually the basal
attachments can be recognized if the calceoli are
lost.
On maxilla 2 medial setae on the inner plate
obviously belonging to the distal fan are discounted
as medial.
NUMBER 245 11
An apicoventral seta or setule on the inner plate
of the maxilliped is discounted in the descriptions
except in the Brolginae where it comes to taxo-
nomic prominence.
Location of measurements for length and width
of articles 5-6 of gnathopods 1-2 and articles
2, 4, 5, 6 of pereopods 3-5 are shown in Figure 5.
Such measurements were taken in mm from the
expanded drawings which for this publication were
reduced to the 44?15 percent level. These measure-
ments are expressed in species descriptions as unre-
duced ratios. The basic measurements in mm are
taken primarily from the holotype but considerable
variation occurs in these dimensions from specimen
to specimen; these data are therefore to be used
only as gross approximations of conditions typical
for the species. Precision in establishing exact
ranges and frequencies of these ratios is left to the
student of population structure.
Counts of setae on article 2 of gnathopods 1-2
and pereopods 1-2 include on the posterior margin
all setae except those in the clump at the postero-
distal corner, but on the anterior margin include all
setae at the antcrodistal corner and on both of the
anterior edges where two such edges are present.
Other counts of setae and spines on pereopods 1-2
refer to facial setae on articles 4 and 5, posterior
spines on article 6 and the presence or absence of
thick posteroproximal spines on article 5. The state-
ment "facial setae on article 4 of pereopods 1-2 = 5
and 6" refers to the presence of 5 setae on article
4 of pereopod I, and 6 setae on article 4 of pereo-
pod 2. If a single number is quoted, each pereopod
bears the stated number. Counts of spines on article
6 refer to the lateral row first and the medial row
second. In the example, "posterior spines of article
6 on pereopods 1-2 = 5 + 6 and 5 + 7 plus mid-
distal seta," each pereopod has 5 lateral spines but
pereopod 1 has 6 medial and pereopod 2 has 7
medial spines. The middistal seta is set between the
rows and occurs on both sets of pereopods.
Facial ridge formulas on article 2 of pereopods
3-5 occasionally have the letter "s" included to de-
note a shortened ridge, for example, the formulas
might read 1 + 1, 1 + 0, and Is + 0 for pereopods
3, 4, 5.
Counts of posterior setae on epimera disregard
the dorsalmost setule or seta, if disjunct; in such
case that setule is described in a subsequent phrase.
The length of article 2 on the outer ramus of
uropod 3, is cited in the descriptions as a decimal,
being the calculation of the length of article 2 as
a fraction of the length of article 1.
NOMENCLATURE.?Except where noted, all new
names derive from the aboriginal languages of Aus-
tralia. All new generic names, regardless of ending,
are masculine in gender. Those Australian species
names not agreeing in ending with their genera are
nouns in apposition. Names are derived from ap-
propriate adjectives or legendary descriptives of
gods, star clusters, biological analogues, or natural
phenomena.
PRESENTATION OF SPECIES DATA.?The description
of a species generally follows a standardized form
established for this study, although occasional de-
viations occur in order to accomodate unusual
morphological conditions.
"Description of Female" (or "Male" if the female
is unknown, poorly known, or included in our col-
lections in subadult condition only) is based on the
holotype and other adults, usually restricted to 10
or fewer individuals. Such specimens, supplemen-
tary to the holotype, are listed under "Voucher
Material." As nearly as possible, descriptions of
species within any genus consider the same set of
characters and use the same terminology and order
to facilitate congruency. From genus to genus, how-
ever, slight to great differences do occur in species
descriptions owing to different taxonomic states.
A female has been selected as holotype wherever
possible. Females of Phoxocephalidae provide a
better taxonomic standard than males because
males have not been uniformly collected and be-
cause males undergo numerous transformations of
characters in adulthood. Small juveniles of several
species, except those from the 1965 (February-
April) survey, are largely uncollected. The smallest
of these are lost through 1 mm sieves. Most non-
adult material in many species is composed of large
juveniles or specimens called "subadults," within
one or two instars of reaching sexual maturity.
These subadults take the form of females.
Little attention has been accorded to variation in
spine and setae counts on gnathopods, pereopods,
and most mouthparts, but antennae, epimera, uro-
pods, and telson of all voucher specimens have been
examined for spine and setae counts or other gross
variables. Details of minor characters are confined
to statements about either the holotype or a similar
adult. For example, setation count on article 2 of
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
the first four legs is usually restricted to observa-
tions made on one specimen. Such constraint to a
narrow typological limit is not intended to be
definitive, but should be sufficient to give some in-
dication of interspecific distinctions that may, in
the future, be worth statistical study.
In addition to noting minor variations within the
voucher material, the section "Observations" con-
tains miscellaneous information and notes on spe-
cial or uncommon features not included in the de-
scription, any of which may be of potential concern
to the validity of the taxonomic appraisals. Unless
otherwise specified, comments in this section refer
to the species as a whole, not just to individual
specimens.
Unusual deviations found in voucher and non-
voucher materials are discussed under "Variations."
Where females are used as the basis of a specific
description, the attendant "Description of Male" is
variable in form and content. Males undergo sev-
eral instars of transformation from the femalelike
morphology of the subadult to the terminal male
stage. Male distinctions are therefore more or less
strong depending on the degree of this transforma-
tion. Fully terminal males of several species have
not been found. Details of transformation are re-
served for a more intensive project studying life
histories of amphipods.
In previous studies such as that of J. L. Barnard
(1960) in the eastern Pacific Ocean, males were
rarely found in bottom grabs but frequently were
caught in neritic tows or near night lights strung
from ships in shallow water. In contrast, males have
been caught abundantly in the bottom grabs from
southern Australia. However, a few neritic tows
have produced males of certain species not found
in benthic samples.
A "Standard Description of Male Phoxocepha-
lids," applicable especially to Birubius, the domi-
nant Australian genus, is presented on page 26. The
details of this standard description are omitted
from the specific descriptions of males unless war-
ranted by deviations from the pattern. Occasional
setal counts on various appendages are cited to
demonstrate similarities to or distinctions from the
female, especially in cases where good male ma-
terial is available. Because the description of the
male is included for the purpose of providing a
contrast with the female and because significant
distinction between the sexes may vary from species
to species, male attributes lacking distinction from
the females are omitted and the format is adjusted
to accomodate specific needs.
Where necessary, a section under the heading
"Illustrations" is included to explain anomalies in
the graphic presentation. If an illustration of any
particular part is omitted, reference may be made
to a comparable illustration for another species; or,
if such illustration is inadequate to portray special
variations or peculiarities, these are described under
"Observations" or "Variations." Sample localities
for illustrated specimens have not been listed in the
figure legends since this information may be de-
rived by correlating the locality designations found
under "Type-Locality" and "Voucher Material"
with the data included in the "Appendix," supple-
mented by Figures 2-4. Details of "Illustration
Technique" are presented below.
Under "Voucher Material" will be found a list of
the specimens, in addition to the holotype, used for
illustration and for confirmatory observations, e.g.,
occasionally an array of variously sized adults is
used to check variation in spine and setae counts.
These specimens, designated in illustrations by
lower case letter, usually have been dissected com-
pletely. The parts are preserved in small vials and
deposited in the National Museum of Victoria or
in the museum from which they were borrowed.
Affinities of the species are discussed in "Rela-
tionship." The discussions are far from exhaustive.
The strongest information concerning one or a few
closely similar species is presented but replications
from species to species are omitted. In the most
complicated section (the 37 species of Birubius),
keys, discussions, and an evolutionary chart assist in
the comparisons; the members of that genus are
arranged loosely in a phyletic order. Many aspects
of growth are poorly understood and in each com-
parison between species the differences have been
limited to those of highest possible qualitative
value. To the reader is left the task of comparing
illustrations and descriptions to find distinctions
that have been ignored for reasons of possible
variability but which in the future may be useful
to the student of populations and thereby ulti-
mately to the alpha taxonomist.
The section under "Material" contains a synop-
sis of identified material using the abbreviations
defined above under "Text Abbreviations."
ILLUSTRATION TECHNIQUE.?The following ma-
NUMBER 245
nipulations have been undertaken to present reas-
onably consistent views of morphology. Generally,
unless otherwise specified in the following para-
graphs or in the labels appearing on drawings,
illustrations herein present a left lateral view.
Head: The lateral aspect is composed with the
eyes and rostral edges in line side to side. Generally,
the anteroventral part of the head is in full view
with the basal article of antenna 2 shown flush to
the side. The dorsal view is presented with the
rostrum fully extended and flush dorsally. Setae and
spines are omitted from antennae attached to the
head. Antenna 1 is viewed with the so-called ventral
margin projecting laterally. Antenna 2 normally
extends outward towards the observer but it is
bent downward in the cephalic view to show the
full width of the articular faces.
Antennae: Antenna 1 is so attached to the head
that the margin termed "ventral" projects more or
less laterally, the dorsal margin turned medially.
In the view of the excised antenna the appendage
is rolled into a position so that the broadest lateral
faces of articles 1-3 are shown to the viewer. On
the excised antenna 2 the same manipulation is
performed but without crushing articles 1-2 before
they are illustrated. Occasionally articles 1-2 are
omitted from the illustrations. Later pressing of the
cover slip presents the full face of articles 4-5.
Prebuccal Complex: The upper lip and epi-
stome are the most unsatisfactory parts for illustra-
tion. Fully facial anterior views are obtained with-
out distortion by frontally orienting the head with
attached prebuccal complex; but the complex is
soft and distortable and the ventral margin of the
upper lip changes appearance with the slightest
change in orientation. A frontal view may show a
slight ventral concavity, an oblique dorsal view
may show a strong concavity, whereas various other
oblique views show truncation or convexity and
may bring into focus a small midhump. Certain
species lack humps and bear concavities in any view
so that we have abandoned reliance on prebuccal
anterior views for important specific distinctions.
Although no Australian taxa, except Booranus,
have the produced epistomal cusp so common to
species in the eastern Pacific Ocean, the prebuccal
complex is illustrated for holotypes of most species
to show subtle differences in conformation.
Mandible: The palpar hump is described as
absent, small, medium, or large in many species
without specifying its precise dimensions. Such
measurements can be learned by examining the
variations depicted in several species. The degree
of bulbosity in mandibular molars may vary ac-
cording to preservational techniques; thus extensive
shadow illustration of this feature is deemphasized.
Mandibular spines are shown in the illustrations as
pressed flat in unnatural orientation; this flattening
may cause the spines to fall in a variety of planes,
thus resulting in their orientation being different
from drawing to drawing.
Lower Lip: This is another appendage that is
difficult to illustrate because it often preserves
poorly, is difficult to orient, and requires the most
tedious of manipulations. The lower lip is difficult
to excise in phoxocephalids without breaking con-
nections closely appressed to the first maxillae and
mandibles. The bundle of parts containing the
lower lip and the pair of first maxillae is excised
and mounted on a slide with the coverslip initially
elevated by sand grains and then lowered so as to
flatten the mandibular lobes of the lower lip with-
out crushing other parts. The oral view of the lower
lip is outlined first, deep focusing is employed to
obtain aboral structures, then the maxillae are
teased away from the base of the lower lip, the lat-
ter turned over, and additional configuration at the
base of the inner lobes is added to the illustration,
the entire view being converted to an aboral aspect.
Maxilla 1: The appendage is first flattened suf-
ficiently to give a view of the flattest face of the
inner plate, which is illustrated aborally. Later the
coverslip is pressed first to flatten the outer plate
and again to press the palp as flat as possible; how-
ever, the palp is rarely flattened to its full width
and thus is drawn in oblique view to avoid crush-
ing of its base and apex. The inner plate often
overlaps the outer plate so greatly that in the illus-
tration the inner plate is moved medially so as to
prevent confusion from overlapping lines.
Maxilliped: A coverslip is mounted lightly over
the appendage to flatten the inner plate first and,
after that is illustrated orally, subsequent pressings
of the coverslip are used to flatten the outer plate
and then progressively the segments of the palp.
Only half of the appendage complex is shown.
Gnathopods (legs 1-2): The medial view of the
right members is drawn preferentially but lateral
views of left members are illustrated if opposite
sides are in poor condition; minor variations in
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
setation on the lateral and medial sides of these
appendages have been ignored. Article 2 of these
appendages is not illustrated, although setal counts
on anterior and posterior margins are given in the
descriptions.
Pereopods 1-2 (legs 5-4): The main taxonomic
features are found on the outer or lateral surfaces
of these appendages. Article 2 is not illustrated al-
though setal counts are cited in the text. Setal and
spine counts of articles 4-6 are compared in the
descriptions. Pereopod 2 is usually similar to
pereopod 1 though often very slightly stouter.
Pereopods 3-5 (legs 5-7): Although the main
taxonomic features are found on the lateral sur-
faces, the medial apex of article 6 on pereopod 5
merits description because of specific variables in
the degree of terminal digitation or in the develop-
ment of fringe and combs.
Epimera: The carcass of the amphipod is rolled
so that the epimera are shown as flush laterally as
possible, although epimera 1-2 roll inward slightly
along the ventral margin. In identifying phoxo-
cephalids one must scan the epimera under high
power microscopy to detect any missing setae by
finding the sockets retained in the cuticle.
Uropods and Telson: Uropods 1-2 are illus-
trated as attached to the animal with supporting
views of medial margins in many species where
spine sizes and counts on medial margins have com-
parative value interspecifically. In preserved phoxo-
cephalids uropod 3 is usually thrust anterodorsad
against the telson; this is the relationship shown in
the lateral view of the pleon and attached struc-
tures. Uropod 3 and the telson are better viewed
from the dorsal aspect after dissection. In making
the drawing of the exised uropod 3 the appendage
is rotated so that the widest flattening of the rami
occurs consistent with avoiding any crushing of the
peduncle. Either right or left uropod 3 may be
illustrated. The telson is especially difficult to
mount and flatten in phoxocephalids. Direct exci-
sion from pleonite 6 is almost impossible because
the telson is usually cleft more than 90 percent to
the base. Therefore, pleonite 6 is removed to a
slide, the tubular pleonite broken ventrally, the
sides spread outward fan-wise, and then teased
away from the telson, especially at the basolateral
corners; unless this is done the telson will not flat-
ten basally.
Cuticle: The elements of the cuticle are so small
that oil-immersion microscopy scarcely resolves
their structure. Sufficient items of taxonomic inter-
est have been found nevertheless, to present a brief
analysis of cuticle for each species. Perhaps the most
favorable locality for viewing is the posterior lobe
of coxa 5 (the coxa of pereopod 3, leg 5). This part
is superior to epimera which are difficult to flatten
without destructive excision and superior to large
flat coxae such as coxa 4 because coxa 5 generally
has all of the gross variables one might encounter.
In many species the epimera lack bulbar setules
and in many other species coxae 1-4 lack variants
of bulbar setules such as pipes and may also lack
striations or have polygonal structures atypirally
formed. Environmental abrasion is reduced on coxa
5 posteriorly.
Reconstructions have sometimes been necessary
because of faulty material; the most frequent of
these involves the edges of coxae and second articles
of pereopods 3-5 which in death and preservation
often shrivel microscopically to the extent that
corners have sharp points, or develop notches in
specimens that were about to molt before death and
in which the cuticle has pulled away slightly from
the subcuticular flesh. These extensions and in-
vaginations have been smoothed over in the illus-
trations. Setae and spines often have been altered
to present a clear picture of their relationship;
their directions of pointing have been moved, not
their origins or insertions. Armaments missing on
appendages or epimera have occasionally been re-
constructed on the illustration by observation of
conditions on opposite side of the specimen. Rare
specimens, especially natatory males, in which the
dorsal part of the eyes is depressed by preserva-
tional defect (dissolution of fatty supportive tissue
by alcohol), have been reconstructed in the lateral
view so as to inflate or move dorsally the mass of
ommatidia into what we believe is a natural posi-
tion; this defect does not show from the dorsal view
because lateral shrinkage evidently does not accom-
pany the dorsal collapse.
Enlargements: As normally practiced in am-
phipodan taxonomy illustrations are given without
scales of enlargements. Terminal adults do not
necessarily have a fixed size and may vary with
season and environmental conditions. The uni-
formity of natatory males suggests to us that they
may have a fixed size but this would be exceptional
in Amphipoda.
NUMBER 245 15
Drawings of specific parts are magnified in pro-
portion to other parts as follows: pereopods 3-5 are
drawn at the same magnification; gnathopods 1-2
are enlarged to a common but unique degree;
pereopods 1-2 are drawn at a common magnifica-
tion, one which is greater than that for pereopods
3-5, in order to show details of articles 5 and 6;
coxae 1-4 are drawn to a scale common to their
group but distinct from those of the various ap-
pendage groups; maxilla 1 is more highly magni-
fied than is maxilla 2; antenna 2 is usually en-
larged to a greater degree than is antenna 1;
mandibular bodies are drawn at larger or smaller
magnifications than are palps depending on details
to be shown; telsons are enlarged more than is
uropod S; head and pleon are drawn at distinct
magnifications; when separate from the urosome,
uropod 2 is enlarged more than is uropod 1; cuticle
is enlarged to the maximum degree with oil-
immersion lens (X 1200) and then reduced in the
figures herein presented to a level of 45 percent of
the magnified image. All other parts are enlarged
sufficiently to portray taxonomic details.
Ink Renderings: All structures on the views of
cuticle have been inked in solid lines despite the
fact that the bulbs and pipes are below the chitin;
the chitin, however, is so clear that these structures
appear suspended in space as in the ink. rendering.
The bases of many spines and setae are inked solid
even though they are inserted into appendages as
shown by their extended bases. Dashed or dotted
lines indicate structures lying behind or below the
main appendage.
In the figure(s) illustrating each species, the chief
specimen depicted is identified first in the figure
legend. Any illustrated subsidiary specimens are
assigned lower case letters that (1) are identified
with the proper specimen in the legend and (2)
appear as components on the left side of the labels
on the drawings of the part reproduced from such
specimens.
Taxonomic Characters of Phoxocephalidae
Phoxocephalidae are among the most richly
diverse of amphipod groups in taxonomic char-
acters. Almost all external parts of the body, includ-
ing head, mouthparts, gnathopods, pereopods uro-
pods, telson, epimera, and coxae, bear characters of
both specific and generic value. The only fully stable
conditions in the family are: the relationship
between the general form and relative sizes of
pereopods 4 and 5; the subchelateness or chelateness
of the gnathopods; and the maintenance of the fol-
lowing parts (often lost in other families): inner
ramus of uropod 3, strong cleft on the telson, dactyl
on the maxilliped, palps on the maxilliped, multi-
articulate accessory flagellum, inner lobes and man-
dibular lobes on the lower lip, and rami on
uropods 1-2 (although inner rami occasionally are
reduced or fused to the peduncle).
The following characters typical of conceptually
primitive phoxocephalids may be obscured by re-
duction or loss: rostrum, flagellum of antenna 2,
thickness of articles and degree of fossorial setation
on pereopods, setation or spination of epimera and
uropods 1-2, presence of article 2 on outer ramus of
uropod 3, size of uropod 3, and elongation of the
flagellum on male antenna 2 (sexual character).
Numerous valuable characters in Phoxocephalidae
have previously been ignored or underemphasized.
Until this study the primary character alternatives
in phoxocephalidan taxonomy were: (1) tritura-
tiveness or simplicity of mandibular molar; {2)
presence or absence of a large palpar hump on the
mandible; (3) presence or absence of an ensiform
process on article 2 of antenna 2; (4) presence or
absence of eyes; (5) bi- or uniarticulation of palp
on maxilla 1; (6) presence of a process on article 3
of maxillipedal palp; (7) proportions of the max-
illipedal dactyl and its apical nail; (8) enlargement
and eusirid (predatorial) adaptation of gnathopod
1; (9) thickness of article 2 on pereopod 3; and (10)
presence or absence of apicodorsal spines on various
rami of uropods 1-2 (J. L. Barnard, 1960, 1969a).
In earlier days the size of article 4 on pereopod
4 and the similarity or dissimilarity of gnathopods
were characters also used in phoxocephalid taxon-
omy (Stebbing, 1906). Size of article 4 on pereopod
4 as a useful character was deemphasized (J. L.
Barnard, 1960) and we have found that this re-
mains true. Such a character alternative signals a
potential generic distinction between two species
but better characters are available to rectify the
generic diagnoses. The dorsal shape of the rostrum
has occasionally been used as a generic character
(Bousfield, 1973) but our studies indicate that the
usefulness is vagarious.
Barnard and Drummond (1976) presented a list
of characters newly utilized in the study of Austra-
16
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
lian phoxocephalids. At least 250 characters in each
species have been examined and assessed in the
Australian study and many of these are highly use-
ful at specific and generic levels of classification.
Many are characters not heretofore widely used
even in many other families, for example: fine
structure of mandibular molars, setation patterns
on mandibular palps, proportions of the setal dis-
tribution on article 2 of antenna 1 and article 5 of
antenna 2, facial spine and setal distributions on
articles 3-4 of antenna 2, presence or absence of cones
on the lower lip, setation counts on the inner plate
of maxilla 1, degree of cleft between inner lobes of
the maxillipeds, linear or nonlinear arrangement of
facial setae on article 3 of the maxillipedal palp,
presence or absence of posteroproximal setae on
article 5 of pereopods 1-2, presence and si/e of the
midapical subdactylar spine on article 6 of pereo-
pods 1-2, apical ornamentation of dactyls on pereo-
pods 1-2, size and shape of article 3 on pereopod 5,
presence or absence of facial setae on article 2 of
pereopod 5, presence or absence of combs in various
places on pereopods and uropods including medial
surfaces and rami, presence or absence of long
flexible setae on epimera and uropods (at generic
level), numbers of and shape of apical setae on
article 2 of the outer ramus on uropod 3, presence
and degree of development of thick spines on
uropods and urosomal surfaces, lengths of rami on
uropods 1-2 with special attention to inner ramus
of uropod 2, size and position of glandular tissue
in urosome, and the presence and number of apical,
dorsal, and lateral spines on the telson.
The shape of prebuccal parts was used at specific
level by J. L. Barnard (1960); subsequently, generic
characters have been discovered in those parts. We
continue to find great value in the shape and orna-
ments of the maxillipedal dactyl, the size of the
palpar hump on the mandible, and general aspects
of gnathopods, but we have also found classificatory
value in the finer structure of gnathopods in terms
of anterior setation on article 6, degree of posterior
concealment of article 5 on gnathopod 2, and
minute difference in size between the pairs of
gnathopods. One subfamily, Tipimeginae, is charac-
terized by minuteness of pereopod 5 and within
this subfamily, shape and ornamentation of epi-
meron 3 have generic value.
Convergence in phoxocephalids is rampant. This
has led to a very unsatisfactory state of classification
in phoxocephalids of the northern hemisphere (J.
L. Barnard, 1960) where convergence has occurred
in what are primarily highly advanced phoxocepha-
lids with apparent origins from the more primitive
stock in the southern hemisphere. An overview of
those from Australia has revealed what appears to
be a more satisfactory solution to the interrelation-
ships of world phoxocephalids, a solution that relies
on the importance of what were considered, hereto-
fore, to be minor characteristics. For example, ge-
neric value is found in anterior setation on the hands
of gnathopods, the presence or absence of an en-
larged inner peduncular spine on uropod 1. and
the presence or absence of supernumerary setation
or spination on the telson. Even more importantly,
the ancestry of Paraphoxu.% oculatus, ihe type
species of a genus whose name has lx*cn used lo em-
brace a host of phoxotcphalids now seen to IK* mis-
classified, now can In- traced.
The classificatory difficulty is well illustrated by
demonstrating the convergence between I'mn/ilioxiis
oculatus and a sympatric species, "P." spinosus
from the northern hemisphere. No valuable char-
acters to distinguish them generically were found
by J. L. Barnard (1960) but study of Australian
phoxocephalids demonstrates that Paraphoxus ocu-
latus is the highly evolved product of the Brolginac
so abundant in the southern hemisphere. "Para-
phoxus" spinosus must be transferred to a new
genus which will be distinguished on the following
slim characters: presence of more than 3 spines on
the molars, presence of 3 (not 2) setae on the inner
plate of maxilla 1, presence of 3 setae on epimeron
3, presence of two full rows of spines on article 4
of antenna 2, full enlargement of the apicomedial
spine on the peduncle of uropod 1, poor develop-
ment of the apical nail on the maxillipedal dactyl,
and the ordinary elongation of article 5 on gnatho-
pod 2. Restudy of "P." spinosus has not been com-
pleted but it would appear to be a very advanced
product of the Parharpiniinae on which telsonic
and molarial setation has decreased but which is
distinguished from the Brolginae in having more
than 3 molarial spines and more than 2 setae on
the inner plate of maxilla 1.
Other examples of convergence and classificatory
difficulties will be reserved for a later paper on
reevaluation of northern phoxocephalids.
EVALUATION OF FEMALES.?An evaluation of taxo-
nomic characters in Phoxocephalidae, as based on
NUMBER 245 17
the Australian fauna and on a survey of the taxo-
nomic literature, is presented below in morpholog-
ical order from head to telson. The same order is
employed under "Description of Females" found in
the presentation of individual species.
Head: The rostrum is elongate, untapered, and
flattened dorsally to form a visor in primitive
phoxocephalids. Overall, the head is more elongate
than in most nonphoxocephalid amphipods but
this is not necessarily a condition correlated with
fossorial behavior because, on the one hand, non-
fossorial amphipods such as ampeliscids have an
elongate head and, on the other hand, in many
genera of other fossorial amphipods in the Gam-
maridae (sensu lato), Oedicerotidae, and Synopiidae
the head is not necessarily elongate. Despite these
exceptions noted, the rostrum may have a fossorial
function in many sprues.
In subfamilies such as Brolginae, Phoxocephali-
nae, and Harpiniinae the head is maintained in
unaltered degree; but in several subfamilies such
as Birubiinae and Joubinellinae the rostrum be-
comes narrowed from side to side, often becomes
shortened, and occasionally becomes obsolescent.
To some extent the most grossly reduced rostra are
associated with other adaptations suggesting com-
mensal istic behavior or functions apart from bur-
rowing {Kondoleus). These adaptations include
nonskid cuticle, reduced flagela of antennae, pred-
atorial gnathopods, pygidization (reduction in
complexity or increase in rigidification of urosomal
structure), development of raptorial spines on
pereopods 1-2, or marked change in structure of
spines on antennae and uropods. Such modifications
may be found in Kotla, Yammacoona, and Japara.
But in a single genus, Birubius, the rostrum pro-
gresses from the fully developed condition through
a series of evolutionary lines to a condition in B.
wulgaru in which it scarcely exists.
The rostrum alone is not necessarily a good
generic character because of these vargarious evolu-
tionary sequences. The narrowed rostrum was at
one time believed to mark the trichophoxin kind
of evolutionary stock in the old umbrella Para-
phoxus-Pontharpinia-Parharpinia classification but
we now recognize that Trichophoxus itself belongs
in a group entirely distinct from other phoxocepha-
lids and that the species formerly included in that
umbrellar designation belong to several distinctive
subfamilies and many genera.
Eyes: Ocular structures are ommatidial in phoxo-
cephalids. Little is known of taxonomic significance
in eyes at generic level except that Pseudharpinia
is maintained apart from Heterophoxus mainly on
the absence of eyes even though J. L. Barnard
(1966a) found the normally oculate Heterophoxus
oculatus to be blind in depths below 200 m off
California. Eyes are degenerate in Phoxocephalus
holbolli but present and well developed in many
other members of the genus.
Color of ocular pigment in life is poorly known
in phoxocephalids although we have seen several
species in life with deeply black eyes. In preservative
these eyes remain black from distant perspective but
usually appear deep purple under high magnifica-
tion. Most species of the subfamily Phoxocephalinae
have almost perfectly clear eyes in preservative,
thereby suggesting that they have red pigments in
life, as red pigments disappear quickly after preser-
vation. We presume that a modicum of taxonomic
information would be available in freshly preserved
collections (J. L. Barnard, 1970).
Eyes in oculate males become greatly enlarged.
These males often swarm in neritic waters and are
attracted to night lights (Fage, 1932, 1933). We
know of no anoculate and benthic deep-sea species
whose males have been caught in epibenthic fishing
and we presume that anoculate males have reduced
swarming behavior.
Antenna 1: Article 1 is elongate and stout and
article 3 is much shortened and narrow in phoxo-
cephalids. Article 2 varies in length but usually
falls into one of two groups, long or short (see
evolutionary scheme of Figure 6). The entire an-
tenna is generally rotated slightly outwards so that
the hypothetically primitive lateral surface faces
somewhat dorsally; this becomes more pronounced
in those species with very broad bodies and heads
and small rostra and is similar to the condition in
the broad-bodied haustoriids. Penicillate setules are
well developed on the ventral margin of article 1
and become very dense in adult males. The dorso-
distal apex of article 1 is often slightly produced
but has no value in taxonomy in the present scheme
because the degree of extension would have to be
measured by micrometrical means. The ventral
setae on article 2 are of great generic value. They
occur in two gross variations, either widely spread
in a crescent ventrally or confined apically. In most
cases the apical confinement is restricted to genera
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
with shortened article 2. Occasional specific differ-
entiation is made between setae that are widely
spread and those confined to the proximal third of
the article. In a few taxa the setae are closely con-
tinuous or severely reduced in number.
The primary and accessory fiagella are multi-
articulate. The articles may be elongate or very
short but little use has been made of this feature
taxonomically (for an exception see Birubius
maamus herein). Aesthetascs of several sizes may be
found on the distal articles, rarely as far proximally
as article 1, and usually restricted to one aesthetasc
per article. Occasional aberrations in the fiagella of
antenna 1 occur. The males of Elpeddo kaikai and
Wildusi fuegiensis have unusual first antennae re-
sembling those of lysianassids in which article 1 of
the primary flagellum has become swollen and
richly furnished with aesthetascs.
Antenna 2: Article 1 is either simple or bears a
weak to greatly enlarged process. When the process
is present, it and the antenna are termed "ensi-
form." The largest ensiform processes are confined
to a few genera in the subfamily Harpiniinae but
these processes are weakly developed in several
genera of other subfamilies and in undescribed
subfamilies from the eastern Pacific Ocean.
Article 2 is very short, largely hidden from lat-
eral view. It bears a gland cone medioventrally.
Article 3 is relatively stout in phoxocephalids. In
primitive phoxocephalids such as Pontharpinia and
Tipimegus, as in the Ponto-Caspian gammarids, it
bears abundant facial setae, but these are reduced
to 2 in almost all others, and may be absent in sev-
eral species recorded in the literature. In Birubius
the 2 setae are small but the more proximal mem-
ber is usually much the smaller.
Article 4 bears facial spines organized into one or
more rows in all phoxocephalids as far as known,
except Joubinella. In some of the genera of Biru-
biinae and Harpiniinae the spines are mainly con-
fined to a subapical position so that they almost
appear to be marginal rather than facial. The ven-
tral margin of articles 4-5 is often richly furnished
with setae of one or more size groups. In the more
primitive taxa, such as Pontharpinia, the long setae
are intermingled with a group of short penicillate
setules but in more advanced taxa the penicillate
setules become reduced to a few proximal members.
The apicoventral face may bear one or more thick
spines. The antennal spines become jewel-like
spheres in Kondoleus.
Article 5 is generally shorter and thinner than
article 4 and in most taxa of such groups as Har-
piniinae and Brolginae article 5 is especially small.
Both articles 4 and 5 become enlarged in males but
in groups in which article 5 is small in the female,
the male article 5 does not enlarge as much as in
groups such as the Birubiinae. in which article 5
is large in the female. Thick apicofacial spines often
occur in females but are reduced to setules in males
of many species. Midfacial spines may be present or
absent infragenerically in Birubius but in such
groups as the Tipimeginae the absence of facial
spines is marked as a generic character.
The flagellum in females is usually shorter than
twice the combined length of articles 4?5 of the
peduncle but it becomes elongate and proliferate in
all male phoxocephalids except those of the genus
Harpinia. This may also be true in some taxa of
Pontharpiniinae and Tipimeginae but we have not
confirmed this at present.
Prebuccal Complex: The presence or absence
of sharp, anteriorly projecting tooth on the epistome
has been a useful character in specific identification
of phoxocephalids (J. L. Barnard, 1960). In the
most commonly encountered prebuccal condition
the upper lip and epistome are mostly fused to-
gether with the weakly distinct upper lip domina-
ting the epistome. In several genera, however, such
as Japara, Kondoleus, and Tipimegus, the two parts
are fully articulate and one must presume that the
variables encountered are relatively uniform at
generic level. Tipimegus and Booranus are well dis-
tinguished by differences in prebuccal parts.
Mandible: The body of the mandible is large
but never as heavily modified as in certain haus-
toriids. A few genera, such as Mandibulophoxus,
and Yammacoona, have that part of the mandible
to which the palp is attached strongly extended;
this is known as the palpar hump.
Right and left incisors are generally dissimilar.
The most commonly encountered right incisor is a
flat cutting edge bounded on each end with a blunt
tooth and with a third inner and subapical tooth
partly hidden by the lacinia mobilis. Occasionally
in certain taxa the cutting edge is divided into four
or more teeth (Kondoleus, Tipimegus). The left
incisor commonly is composed of two weakly
divided branches each of which has one or more
additional subdivisions, rarely with a more evenly
NUMBER 245 19
distributed series of teeth.
The right lacinia mobilis has proved to be in-
valuable for specific identifications in this study.
This structure has been ignored in earlier literature
and often depicted so poorly that its character
cannot be determined. The evolutionary deployment
(Figure 82) of Birubius is based on the progressive
simplification of the right lacinia mobilis. The pre-
sumed line of development commences with a
broad, flabellate, and denticulate piece which
becomes clearly divided into two pieces, the distal-
most of which remains flabellate, whereas the prox-
imal branch assumes the appearance of a conical
raker spine. In the next stage the distal branch is
also reduced to a simple conical form, then decreas-
ing in size until, with its loss, the right lacinia
mobilis almost fully resembles an ordinary raker
spine. There may be instances in which the right
lacinia mobilis is actually lost (Birubius eleebanus
and B. wulgaru).
An equally logical supposition would be that the
right lacinia mobilis becomes more complex by the
fusion of one raker spine to a primordial lacinia
mobilis so as to form the intermediate condition
but the decaying progress found in Birubius fits
the advancement of other characters, such as general
loss of setation and reduction in rostrum. The right
lacinia mobilis of Pontharpinia, the primitive
phoxocephalid, appears to be composed of just the
distal flabellate piece, so we hypothesize that the
ancestral Birubius was characterized by the joining
of a raker spine to the lacinia mobilis but that this
new adaptation rapidly was reversed through the
progression we have outlined in Figure 82. In any
event the right lacinia mobilis is highly variable in
other phoxocephalids, sometimes appearing to be a
simple broad flake and in other places appearing to
have a proximal branch that resembles a raker
spine.
The left lacinia mobilis is more stable than the
right in phoxocephalids. It generally consists of a
palmate piece with 4-6 apical teeth, rarely with the
teeth blunted or reduced (Kotla), but often in
Birubius with the middle teeth shortened.
Raker spines are well developed, numerous, and
not generally diverse in phoxocephalids except that
a few proximal rakers are sometimes smaller than
the others, suggesting that rakers are proliferated
basally as growth of the animal proceeds. Between
raker spines there are frequently accessory pluse-
tules, alternating with rakers, but these are lost in
such aberrant clusters as the Birubius babaneekus
group where the rakers themselves become very
short and broad.
In earlier literature only two kinds of molar were
recognized, triturative and non-triturative, but we
now find at least five basic kinds of molars in
phoxocephalids (Figure 5) and one may see from
inspection of our illustrations that further subdivi-
sions may be necessary in the future. Stage one, the
primitive molar (Figure 5: A), we presume to be the
fully triturative form typical of Pontharpinia and
Phoxocephalus and a few other genera. This molar
is not precisely like that found in gammaroid am-
phipods but is a subcylindrical acetabulum-like pro-
jection with a weakly hollowed apex surrounded by
a crown of teeth. The second stage (Figure 5: B) is
represented by Leongathus in which the molar ap-
pears as if only one side of the cylinder and its teeth
have remained. Stage three (Figure 5: C) appears
in the Tipimeginae in which the remaining half of
the molar becomes more conical and retains only
three or four of the teeth, apparently semiarticulate.
This form might be hypothesized to have developed
from either stage two or stage four as the apparent
articulation of the teeth suggests that they could be
enlarged articulate spines. Stage four (Figure 5: D)
is found in most phoxocephalids and comprises a
small conical or pillow shaped protrusion on the
apex of which occur 4 or more splayed and small
semiarticulate spines. One may see from the illustra-
tions that this kind might be subdivided into sev-
eral subgroups based on size, number and plumosity
of the spines and the particular shape of the
molarial body itself. We see major categories of
classification in Birubius alone as "elongate
plaques," "short bulbs," "soft cones," and "molar
absent but position marked by spines alone." The
final stage (Figure 5: ?) is typical of Brolginae in
which only three or fewer semiarticulate spines re-
main. These often become enlarged, or become
joined basally, and occasionally arise from a dis-
tinct basal plate.
We do not suggest that these stages normally oc-
cur in the sequence proposed in every evolutionary
line. They have been arranged simply in an order
of decreasing complexity. Nevertheless we do sug-
gest that the missing links between the fully tritura-
tive and the weakly spinose nontriturative molars
might be represented by stages two and three and
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
which might well be called "semitriturative" and
"pseudotriturative" to reflect their distinctions and
possible functions.
The presence of a 3-articulate mandibular palp
is a stable feature of Phoxocephalidae. Article 1 is
rarely elongate or armed with setae. Article 2 is
elongate, usually bears at least 2 setae, and occa-
sionally, bears facial setae in more primitive taxa.
Article 3 is well developed, setose facially, usually
truncate obliquely, bears stiff apical spine-setae but
only in occasional males are setae developed on the
inner margin (Tipimegus). In primitive phoxoceph-
alids the outer margin bears a notch armed with
one or more setae, but in the more advanced phoxo-
cephalids these setae have shifted onto the face of
article 3 or are lost entirely. They may occur in
units or pairs (or greater counts) on one or both
faces of article 3. The trend appears to be towards
total loss in the more advanced taxa.
Lower Lip: Phoxocephalids have well developed
mandibular lobes and fleshy inner lobes. The inner
lobes are tightly appressed or partially fused to-
gether, with the apices almost truncate or weakly
bulbous. The widely separated outer lobes have
moderately extended mandibular lobes apically
rounded or weakly attenuate. In all but primitive
phoxocephalids each outer lobe has one or a few
salivary cones (extensions of meatal ducts). Thick-
ened setae may also occur in the vicinity of the
cones and some of these may prove to be additional
cones. Inner and outer plates are often minutely
setulose (pubescent) along apical margins.
Maxilla 1: In comparison to primitive gam-
maroids the inner plate in phoxocephalids becomes
enlarged relative to the outer plate but has lost
most of its setae. The apex is usually rounded
broadly and carries either 4 or 2 setae in the major-
ity of phoxocephalids. The primitive Pontharpinia
carries more than 4 shortened setae but Phoxoceph-
alinae have lost all setae on the inner plate and the
surface area is expanded. In Birubius the usual
setation pattern consists of one large inner facial
pluseta, a second often somewhat smaller and more
terminal pluseta, and 2 much smaller apicolateral
setae. In many Brolginae the 2 apicolateral setae
apparently have been lost.
The outer plate bears 5, 7, 9, or 11 spines; in rare
cases an even number of spines occurs (Japara). The
spines are often divided into 2 sets of 5 and 4 or 6
and 5, the smaller set comprising small forked
spines, the larger set comprising large spines, the
medialmost of which are serrate medially, the lat-
eralmost of which are very stout and poorly serrate.
In many genera one of the large lateral spines,
usually spine 2, is especially thickened. In the genus
Japara, the innermost spine is diverted to the
medial margin and points medially. Many of the
Brolginae have an additional medial cusp forming
a false spine lying almost flush, and therefore hid-
den, along the normal medialmost serrate spine.
Variations in fineness of serrations, as in Kuritus,
have specific usefulness in taxonomy.
The palp of maxilla 1 is either uni- or biarticu-
late but it is rarely reduced to any degree. A baso-
lateral wing usually occurs on the acclivity of the
outer plate below the juncture of the palp. Article
2 is much longer than article 1 and is apically and
often facially armed with spines and setae. Facial
setae are especially well developed in Tipimeginac.
There is little difference between the palps of right
and left sides.
Maxilla 2: The pair of plates forming this struc-
ture is relatively uniform. Unlike many members of
Gammaridae, the inner plate lacks an oblique facial
row of setae. In Brolginae and especially in Phoxo-
cephalinae the plates are shortened and apically
rounded and the setation is heavily reduced.
Maxilliped: This is well developed, with only
an occasional taxon showing some reduction in the
size of the plates (Phoxocephalinae, Yammacoona).
The apical armament on the inner plate of the
primitive phoxocephalid appears to consist of
evenly extending elongate setae but in the majority
of phoxocephalids one or more (usually not more
than 3) short and very stout apical spines have ap-
peared and the setation density has decreased. The
trend in the more advanced evolutionary lines such
as Brolginae and Phoxocephalinae is towards a de-
crease in setation and/or narrowing of the inner
and outer plates. In several of the Phoxocepha-
linae the broad and heavily spinose and setose outer
plate typical of the primitive phoxocephalid be-
comes very thin and the armament is reduced to 4
or fewer spines. This is associated with other losses
of fossorial adaptation in those taxa. The richly
setose palp of the primitive phoxocephalid follows
this same pattern of reduced setosity, but the palp
with the most abundant and scattered facial setae
on article 3 occurs in the Tipimeginae. This article
becomes strongly produced apically in the Lepto-
NUMBER 245 21
phoxus group but is more weakly produced in a
variety of other unrelated taxa. The inner plates
of Kondoleus bear remarkable basal salivary spouts.
The primitive dactyl appears to be long, ungui-
form, and armed with a poorly developed apical
nail which in more advanced taxa becomes much
more elongate while the article itself may become
shorter and stubbier. Supernumerary setules de-
crease in number in the advanced taxa though ex-
ceptions do occur in Microphoxus and several of
the harpiniins, where accessory apical setules be-
come almost as elongate as the nail.
Coxae: In amphipods the coxae are a specializa-
tion of the body form to be discussed apart from
their respective legs. Conformation of the coxae in
phoxocephalids is very stable and basic compared
with the extremes of diversity found in many other
families of amphipods. Coxa 1 is generally adz-
shaped and unreduced in size, coxae 2-3 are elon-
gate rectangles, and coxa 4 forms a broad shield
with a posterodorsal excavation bounded by a
posterior tooth.
Only in the rarest cases do these first 4 coxae be-
come heavily modified (Kotla, Kondoleus), though
in a few taxa, such as Birubius booleus and Ticka-
lerus birubi, coxa 1 is rectangular and elongate.
Coxa 4 is more often variable in size and form,
being either large or extremely large (Basuto). The
ordinary coxa 4 has distinct anterior, ventral and
posterior margins below the posterior tooth but in
many cases the posterior and ventral margins merge,
so that the resultant posterior margin becomes
divergent from the tangent of the anterior margin;
and rarely the ventral margin becomes so short that
the coxa appears subacuminate (Birubius nammul-
dus). Anterodorsal humps are especially prominent
in the Tipimeginae. The presence or absence of
ventral setae on coxa 4 is a very useful taxonomic
character in Birubius and in other genera.
Apparently all phoxocephalid coxae 1-4 bear a
small anteroventral setule usually surrounded by a
clear bulbar halo, though occasionally this halo is
reduced or absent as in the Birubius taldeus group.
Coxa 4 almost always bears a second example of
this setule at the posteroventral corner, this occa-
sionally becoming weakly elongate. Only a few
genera, such as Booranus, have additional ventral
setules of extremely small size.
All phoxocephalids have elongate setae on coxae
1-3. These are severely reduced in number in many
of the advanced taxa such as the Phoxocephalinae
but the greatest reduction has been found in the
Birubius taldeus group where coxae 1?3 each have
only 2 long setae. The occurrence of setae on the
posterior margin of coxa 4 beyond the hypothetical
posteroventral corner occurs mainly in the Tipi-
meginae and in Pontharpinia, the most primitive
taxa of the group.
Coxal gills are generally found on pereonites
2-7 but they are absent or reduced on pereonite 7
in Tipimegus and Limnoporeia. In Limnoporeia
they are absent on pereonite 2. Hence the formulas
2-7, 2-6, or 3-6 may prevail. The gill of coxa 7 is
usually the smallest. Brood plates occur on pereon-
ites 2-5 in all genera except Limnoporeia and
Uldanamia where the plate of pereonite 2 is absent
and the formula is 3-5 only.
Gnathopods: The primitive gnathopodal state
in phoxocephalids is presumed to be that reflected
in large degree by Pontharpinia, in which gna-
thopod 2 strongly dominates gnathopod 1. Unlike
the condition prevailing in gammaroids, article 5
of gnathopod 2 in primitive phoxocephalids is so
short that the posterior margin is hidden from con-
tact with the environment by the abutment of
articles 4 and 6. This condition is termed "cryptic
article 5." If this is an obligatory characteristic of
primitive phoxocephalids and if phoxocephalids in-
deed are descendants of gammaroids then the ma-
jority of phoxocephalids have undergone a reversal
in the sense that article 5 becomes more elongate
in many of the more advanced lines of phoxo-
cephalids, especially in the Birubiinae and unnamed
subfamilies of the eastern Pacific Ocean. A close
approximation of the cryptic condition is, however,
maintained in the advanced Phoxocephalinae and
Harpiniinae although one may see that the most
advanced brolgin, Paraphoxus, has lost some of the
degree of masking found in several of the more
primitive Australian brolgins.
The functional morphology of gnathopods, even
in the well studied gammaroids, is very poorly
known. Gnathopodal function is clearly for grasp-
ing, cleaning other appendages, lowering the top
heavy center of gravity of the body at appropriate
times, and manipulating objects by means other
than grasping. Such a prehensile member is useful
in grasping objects of the inert environment, grasp-
ing other biota in a predatorial mode, grasping the
opposite sex during copulatory amplexus, and pos-
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
sibly assisting in the transfer of spermatophores
from male to female. Among numerous other more
finely characterized functions, shoving capabilities
could be envisioned that include defense or moving
eggs about in the brood pouch.
We hold to the belief that the first true amphipod
had gnathopod 2 fully developed as a grasping de-
vice but that the general trend in many evolution-
ary lines is the reduction in size and prehensility of
that appendage with comcomitant loss or readapta-
tion of many of the grasping functions found in the
more primitive amphipods. We theorize that the
most primitive amphipods had both gnathopods
1-2 of equal size and prehensile capability but that
the usefulness of gnathopod 1 in tandem with
gnathopod 2 was quickly lost by natural selection
in favor of new functions, particularly that of
cleaning anterior appendages and manipulating
small environmental particles. This follows the logic
of accepted peracaridan theory that the maxillipeds,
once fully thoracic, are now mostly cephalic and
that the result of such a trend would be that any
order of crustacean succeeding amphipods would
be one with gnathopod 1 also assuming cephalic
functions. The tendency for free thoracic segment
3 now occasionally to develop a prehensile ap-
pendage is confirmed in such gammai ideans as
Endevoura (Lysianassidae), Parandeniexis (Stego-
cephalidae), and Falklandella (Gammaridae, sensu
lato).
There are many examples, of course, where en-
largement of gnathopod 1 to a condition dominant
over gnathopod 2 has occurred, but we consider
these as "derived" evolutionary rather than as
examples of primitive conditions in which gnatho-
pod 2 is undergoing an evolutionary enlargement
to catch up with gnathopod 1. The aorid examples
are obviously instances in which an enlarged
gnathopod 1 has selective advantage to the tube
dwellers. The pelagic predatorial examples of
Joubinella and Trischizostoma are clearly cases
where the predatorial mode is well served by ex-
tensible and raptorial first gnathopods. In the cases
with which we are most concerned, such as certain
species of Gammarus and Anisogammarus in which
gnathopod 1 is only subtly larger than gnathopod
2, we suggest that the distinctions in function are
also very subtle and that gnathopod 2 is not simply
adapting slowly to match gnathopod 1. Most of
these gammarid taxa have gnathopods complexly
ornamented and spined to such a degree as to be
incomparable to the more simplified and probably
more primitive predatorial kind of gnathopod seen
in the Baikalian acanthogammarids (Dybowsky,
1871). The selective adaptability of gnaihopods
would appear to be rapid and extensive in an evo-
lutionary sense as noted by J. L. Barnard (1970),
for example, the axial reversal of gnathopods in
certain corophiid amphipods.
If one therefore assumes that Joubinella is simply
a predatorially adapted case, then one may state
that phoxocephalids are well advanced over their
probable gammaroid ancestors in respect to gnatho-
pods. Gnathopod 1 is fully modified in shape and
structure to match the ordinary kind of gnathopod
seen in the very advanced gaminaridean families
and genera such as the nonaorid corophiids, tali-
troideans, and dexaminids. The enlarged and highly
prehensile gnathopod 2 is retained in phoxocepha-
lids in subfamilies otherwise noted for their highly
advanced adaptations such as thin-mem bered
pereopod 3 of Harpiniinae and reduced complexity
of mouthparts in Phoxocephalinae. Gnathopod 2 of
the otherwise ordinary Birubiinae has reached the
most advanced or reduced state, where in many
taxa it has article 5 elongate and in overall form
almost precisely matches gnathopod 1. The severe
initial reduction in article 5 of gnathopod 2 has
been partially reversed in the advanced Phoxo-
cephalinae and Harpiniinae and fully reversed in
the Birubiinae and Tipimeginae. In the latter group
and in unnamed subfamilies of the eastern Pacific
Ocean a further adaptation has been added to the
gnathopods in the form of an apicoanterior brush
of setae and spines on the hand. The palms have
become shorter and more transverse than in other
phoxocephalids. A somewhat similar situation oc-
curs in the limnoporeiin group of Phoxocephalinae
where the thin probe-like gnathopods have become
strongly chela te.
Pereopods 1-2: These appear to be universally
fossorial in the Phoxocephalidae. Article 4 of both
pairs is elongate and often greatly broadened. In
the best adapted fossorial species it is heavily setose
posteriorly but a reduction in setosity does occur,
most notably in advanced species of Birubius where
pereopod 2 may have as few as 5 posterior setae on
article 4.
Article 5 occurs in two main forms in Phoxo-
cephalidae. In the Tipimeginae it is elongate and
NUMBER 245 23
bears only terminal posterior setae whereas in most
other phoxocephalids article 5 is swollen and bears
posterior setae well proximal towards article 4. In
most phoxocephalids one or more of the apicopos-
terior setae on article 5 are heavily thickened as
spines and in several species of Birubius (especially
species 22-28) one or more of the proximal setae
are likewise thickened. Article 5 bears apicomedial
combs in the Tipimeginae.
Article 6 is elongate and thin but the posterior
margin is furnished with two rows, lateral and
medial, of spines or setae (or mixtures of both,
especially in the Brolginae and Phoxocephalinae).
In many advanced taxa, especially those which may
live on extremely fine sediments, these spines are
reduced in prominence, either by a lesser number
or a greater flexibility. Many phoxocephalids also
bear an apical seta or spine near the base of the
dactyl, situated between the two posterior rows.
This seta is severely reduced in such genera as
Birubius and occasionally may be absent, whereas
in the Tipimeginae it is large and often superen-
larged as a thick spine longer than the dactyl.
The primitive dactyl of phoxocephalids is com-
plex in the sense that it bears (1) an apical nail
mostly immersed in all taxa, (2) an inner tooth
forming a crotch in which is inserted a stiff but
sinuous setule, and (3) a medial or outer basal
plusetule. In certain taxa of Tipimeginae the med-
ial setule is lost and in the most advanced brolgins
the inner tooth is lost but the raphe marking the
insertion of the setule is retained.
Pereopods 3-5: All are fossorial and pereopods
4-5 maintain their special relative sizes and shapes
throughout the family although numerous modifi-
cations occur that have taxonomic value. The prim-
itive phoxocephalid apparently was characterized
by extensive fossorial adaptations in breadth and
setosity of pereopods 3-5 but advanced taxa have
many of these severely reduced. Article 2 of the
basic pereopod 3 is broad and of even dimensions
axially, but the Harpiniinae are characterized by a
very thin article 2 scarcely matching the width of
article 3. Article 2 tapers in the Parharpiniinae and
Mandibulophoxus. Posterior and, to some extent,
facial, setosity is well developed on article 2 of
primitive phoxocephalids and is maintained to
some extent in the Parharpiniinae but most odier
groups have lost these features. Article 4 and, more
or less, article 5 of pereopods 3-4 is very broad in
primitive groups but in the more advanced Brolgi-
nae, Parharpiniinae, Phoxocephalinae, and Har-
piniinae the stoutness and spinosity of pereopods
3-4 are greatly diminished. In many cases one
might presume this to be an adaptation to fineness
of sediments especially in the deep-sea Phoxocepha-
linae and Harpiniinae. But, as diminution also
occurs in the Birubiinae from the shallows of Aus-
tralia it may have other adaptive significance.
The hatchet shield of article 2 on pereopod 5 is
consistent throughout Phoxocephalidae and resem-
bles that seen in pontoporeid amphipods. All shal-
low water Australian phoxocephalids have a rela-
tively uniform article 2 in terms of the posterior
teeth, which are very small, actually the size of
serrations, whereas in several northern groups and
deep-sea groups some of these teeth become greatly
enlarged or fully developed as spikecusps. The ven-
tral margin of article 2 apparently was heavily
setose in primitive phoxocephalids (Pontharpinia,
Tipimegus) but these setae are lost or reduced to
the level of microscopic setules in the majority of
phoxocephalids. Weak ventral serrations on the
primitive members become greatly enlarged in cer-
tain advanced harpiniins (Heterophoxus) but are
obliterated in the Phoxocephalinae and are ob-
solescent in the majority of phoxocephalids only to
crop up in several evolutionary lines of Birubius,
possibly an indication of diverse origins of the sev-
eral species groups. Facial setae on article 2 occur
rarely, primarily in Pontharpinia and Tipimegus
as a mark of their primitiveness. In Pontharpinia
the setae occur in bundles as they do in Ponto-
Caspian gammaroids, but in Tipimegus they are
solitary.
Pereopod 5 of the Tipimeginae is dwarfed, a
condition unique to that subfamily. Also, article 3
is greatly enlarged in contrast to all other phoxo-
cephalids except to some extent in the Harpiniinae
and perhaps a few other non-Australian taxa. Arti-
cles 4-5 are somewhat thin in Tipimeginae but in
the odier phoxocephalids are more ovate or trape-
zoidal. Apical combs on articles 5-6 have taxonomic
value in several groups (Tipimeginae for example),
and many species of Birubius have digitate proc-
esses apically on article 6. Several males of Birubius
and males of Japara bear enlarged and oddly
shaped spines on article 5. The dactyl of pereopod
5 is usually well developed and in Pontharpinia
has supernumerary marginal setae, but in Yan and
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Kulgaphoxus the dactyl becomes vestigial.
Epimera: The shapes and armaments of the
plates on the sides of pleonites 1-3 are so important
to classification in this group of Australian species
that we believe almost any adult can be identified
to species by reference to epimera and a few other
supporting characters. The epimera of primitive
phoxocephalids might be envisioned as having
marginal setae not only anteriorly, ventrally, and
posteriorly but facially as well, if Tipimegus and
Pontharpinia together can be considered to repre-
sent the ancestral condition.
Epimera 1-2 of the ordinary phoxocephalid each
bears a facial ridge near the ventral margin but only
in a few phoxocephalids do facial setae appear dor-
sal to that ridge line. They are barely dorsal on
epimeron 2 of Pontharpinia but fully so in many
Tipimeginae. Facial setae are rare on epimeron 3
in species outside of the Australia-New Zealand
area where they occur prevalantly in Ponthar-
piniinae, Tipimeginae. and Birubiinae. In the
Birubiinae the evolutionary trend is towards loss of
both facial and marginal setae on epimera and this
trend is characteristic of all other world Phoxo-
cephalidae even in the Brolginae of Australia, a
group characterized by the loss of all long setae on
epimeron 3. Posterior marginal setae on epimera
1-2 are lost rapidly in most evolutionary sequences
although they occur as far east as North America
and may prove useful in tracing the ancestry of the
American fauna. The crowding forward of facial
setae below the lateral ridge in various species of
Birubius is a useful taxonomic character. In Biru-
bius special attention is devoted to the presence,
absence, and size of a posterodorsal seta or setule
on epimeron 3 in tracing phyletic descent.
Pontharpinia bears a large posteroventral tooth
on epimeron 3. This characteristic has been used
by us as one of the starting points in delineating
the evolution of the species groups in Birubius.
This tooth is reduced or obliterated in most phoxo-
cephalids though it crops up in various harpiniins
and in the Waitangi group of the Tipimeginae.
Urosome: Dorsal cuspidation of the body in
phoxocephalids is rare and confined to the urosome.
Kulgaphoxus and Tickalerus have a sharp dorsal
hook on urosomite 3, which occurs also in Methar-
pinia cornuta Schellenberg. The odd "Paraphoxus"
vigitegus J. L. Barnard (1971) bears a dorsal hook
on the urosome in a position presumed to be on
urosomite 2. In that species and in many species of
Birubius urosomites 1-2 are partially to fully fused
together, a condition not otherwise common in the
family.
Lateral spination on the urosome appears to be
of rare occurrence, having been found only in
Birubius lorus and Trichophoxus capillatus.
Ventral and ventrolateral setal brushes are common
in Tipimeginae and a few other taxa and spines or
setules are often found on urosomite 1 near the
base of uropod 1. Conspicuous urosomal glands are
seen in Kondoleus and Kulgaphoxus and in the
former genus cause the urosome to bulge dorsally;
meatal spouts also occur in Kondoleus.
Uropods 1-2: These are very important in the
generic classification of Phoxocephalidae. Basic uro-
pods 1-2 have an elongate |>edunrle with widely
spread dorsal spinosetation on both lateral and
medial margins. The rami extend almost equally
and are spinosc continuously to the apex.
There is a general trend in advanced phoxo-
cephalids to lose most of the medial and lateral
spination or the peduncles so that the remnants are
confined apit ally. Spines or setae generally become
shorter and thicker. A subapical gap in dorsal
spination of the rami on uropods 1-2 appears early
in the evolutionary scheme but in almost every
subfamily there is found a genus retaining almost
continuous dorsal spination on one or more of the
rami. Some of the Tipimeginae have 2 rows of
dorsal spines on the inner ram us of uropod 1. In
Birubius the remnants of continuous spination are
marked in a few species by the presence of an acces-
sory apical nail on one or more rami; and in the
B. taldeus group this becomes vestigial and is
termed an accessory setule. Apical nails are retained
in many genera but in the most advanced Phoxo-
cephalinae, for example, the apical nails are re-
sorbed or so deeply immersed that they become
almost indiscernible.
The presence of basofacial setae on the peduncle
of uropod 1 is primarily a valuable taxonomic
feature of Birubius. The Tipimeginae are char-
acterized by a ventral spike on the peduncle of
uropod 1.
In Pontharpiniinae, Tipimeginae, Brolginae,
Parharpiniinae, and occasionally elsewhere, one of
the apicomedial spines on the peduncle of uropod
1 becomes greatly enlarged and is shifted onto the
apical margin disjunctly from the true inner mar-
NUMBER 245 25
gin. We call this a "special enlarged spine." It is
one of the main characters distinguishing Parhar-
piniinae from Birubiinae. It has generic value in
the former subfamily because in Protophoxus it
occurs not medially but laterally.
The peduncle of uropod 1 is shortened in Tipi-
megus and the outer ramus of uropod 1 occasion-
ally is shortened in the Phoxocephalinae. The inner
ramus of uropod 2 becomes shortened and ulti-
mately fused to the peduncle in Kotla, Kondoleus,
and Matong. The spines on that uropod become
very jewel-like in Kotla, but jewel-like spines are
also seen in several of the undescribed genera in the
eastern Pacific fauna.
Combs on the rami of uropods 1-2 are an im-
portant specific character in the Tipimeginae. In
Brolginae and Phoxocephalinae apical combs on
the peduncles of uropods 1-2 are important fea-
tures but not universally distributed in those sub-
families.
Uropod 3: This is generally of the variramus
type (Stock, 1971) in which the inner ramus of the
female is shorter than the outer but is not fixed to a
specific si/e ("feminine" form) and in the male the
inner ramus elongates to match the outer ramus
equally or subequally ("masculine" form). The
inner ramus is poorly setose in juveniles and in
adult females of many species though in other
species it may be strongly setose. In males it be-
comes heavily setose. The outer ramus usually bears
a second article and in the female has sets of lateral
spines in acclivities. In the male setae are present
in addition to these spines. In primitive phoxo-
cephalids article 2 of the outer ramus is of the
elongate form, about 25 to 35 percent as long as
article 1. This is generally similar to that of pre-
sumed ancestral gammaroids. In advanced phoxo-
cephalids such as Brolginae and Phoxocephalinae,
setae and spines of uropod 3 become reduced but
article 2 of the outer ramus is maintained in
elongate form, whereas in other groups such as
Birubiinae the setae are maintained but article 2
of the outer ramus becomes reduced in size. In
genera such as Japara and male Matong, the second
article is lost and in Kondoleus, Kotla, and Kulga-
phoxus both rami of uropod 3 are also reduced to a
size approximating the length of the peduncle.
Apical setae on the outer ramus are reduced from
the normal two to one in Brolgus but are increased
to three in the Tipimeginae.
Telson: The general shape, size, and degree of
cleavage on the telson remain fairly stable in
Phoxocephalidae though in some of the more ad-
vanced groups such as Phoxocephalinae the telson
becomes elongate. The spination patterns, however,
have taxonomic significance at generic level. The
ordinary phoxocephalid telson bears very few
apical spines and setules and a pair of dorsolateral
setules on each side, but the primitive phoxoceph-
alid telson as represented by Pontharpiniinae,
Tipimeginae, and Parharpiniinae and unnamed
subfamilies of the eastern Pacific, has supernumer-
ary dorsal and lateral spines or setae as are found
in many of the presumed taxa of the ancestral
gammaroid stock. Pontharpinia bears a large brush
of dorsobasal setae on each side of the telson and
the Parharpiniinae have extra dorsal setae or spines
distinguishing them from the Birubiinae. The ordi-
nary male phoxocephalid often has a single longi-
tudinal row of tiny dorsal denticles on each lobe of
the telson but primitive phoxocephalids in the
Tipimeginae have these in broad patches. Some
taxa have the dorsolateral setules shifted apically
(Japara).
Cuticle: The cuticle of ordinary phoxocephalids
usually bears bulbar setules with moderately emer-
gent bristles, often plumose, divided, or of ragged
form. The actual surface is often faintly striate in
the form of human fingerprint patterns (but not
whorled); these striations have been examined with
SEM and found to be the edges of long rows of
overlapping scales. In various species of Birubius,
Matong, and Kotla the cuticle becomes more com-
plex by development of plaques around the bulbar
setules. Parharpinia villosa has narrow pyramidal
irruptions of the cuticle visible at low magnifica-
tion. The ultimate non-skid cuticle is reached in
such genera as Kondoleus where polygons are
formed by an increasing density of overlapping
scales to such an extent that they can be seen under
low power microscopy and can be palpated by
human touch.
Well developed non-skid cuticle is associated
both with species of clear fossorial mode (Parhar-
pinia villosa) and with taxa bearing other morpho-
logical advancements suggesting adaptation to an
inquilinous mode (Kondoleus).
Undoubtedly, electron microscopy will reveal a
host of taxonomically significant structures in cuti-
cles of amphipods and we trust that this will be-
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
come a routine procedure in the near future. In the
views of cuticles we have taken, sufficient variation
occurs, between the sexes of a species and between
individuals of a species, to suggest that problems of
age and elapsed time within an instar may be fac-
tors to be considered in establishing the precise
cuticular configuration. Proliferation of rudi-
mentary bulbar setules may occur at or near sexual
maturity, and at, near, or immediately after ecdysis;
cuticular striations may be lost owing to edema
just before ecdysis; scales may be best developed
just prior to ecdysis; and ultimate body size may
be positively associated with numbers of bulbar
setules.
The ordinary bulbar setule has an emergent
plumose blade of length about equal to or slightly
greater than the double follicular bulb from which
Standard Description of Male Phoxocephalid
(Mainly Birubius)
Body slighter or smaller than in female;
eyes greatly enlarged; article 1 of antenna 1
with medial patch of fuzz, ventral margin
with additional setules; articles 3-4 of antenna
2 with dorsomedial fuzz, facial setae on article
3 shortened, ventral setae of articles 4-5
shorter and fewer than in female, article 5
and flagellum of antenna 2 elongate, furnished
with calceoli; article 3 of mandibular palp
with increased number of basofacial setae;
setae on gnathopods and pereopods shorter
and fewer than in female; article 2 of pereo-
pod 5 and usually of pereopods 3-4 narrower
than in female; epimera 1-3 usually broad-
ened, posterior margins of epimera 1-2 pro-
truding, posterior margin of epimeron 3 short-
ened, setae often fewer; spines of uropods
1-2 smaller than in female, usually more
numerous on peduncle of uropod 2, rami of
uropod 3 "masculine," inner ramus elongate
and setose on both margins, outer ramus with
setae in addition to lateral spinosity of female;
telson elongate, apical spines shortened, each
lobe frequently with dorsal row of denticles;
urosome smaller than in female; pleopods
enlarged, peduncles powerful, peduncle of
pleopod 3 conspicuously shortened.
it grows, but length of the setule varies from species
to species and from bulb to bulb on the same
specimen. Several species have anomalous setules,
branched or extremely short and scarcely emergent.
MALE PHOXOCEPHALIDS.?Terminal male phoxo-
cephalids differ from females in numerous sec-
ondary sexual characteristics. In many cases prim-
ary taxonomic characters also are modified so that
the proper recognition of the two sexes of a species
by taxonomists is occasionally obfuscated. Fully
terminal males of several species are undescribed
because samples of phoxocephalids obtained in bot-
tom grabs often lack associated males of advanced
age. Males apparently swim into the nekton of
neritic waters, or become epibemhic, or are such
powerful swimmers that they can escape grab de-
vices descending towards the bottom. Neritic net
hauls at night often contain male phoxocephalids
but rarely females.
Subadult males and "juvenile males" are com-
monly found in the benthos. "Juvenile males" show
the first external signs of sexual differentiation in a
slight lengthening, due to proliferation of proximal
articles, of the flagellum of the second antenna,
and a slight increase in the size of the eyes. Apart
from this, young males resemble the female, but the
subadult males of some species are noticeably
heavier bodied and rather larger than adult females
of the same species from the same sample. Body
conformation also changes greatly in the later in-
stars, resulting, as a rule, in adult males of slighter
build than the females. Details of transformation
should be reserved for a more intensive project on
the life histories of amphipods.
The following morphological changes occur in
males, but not all in any one taxon. Among those
listed below are some believed to occur so con-
sistently in male phoxocephalids that they have
been incorporated into a standard male description
(above) and then omitted from the specific descrip-
tions of the males in this study:
1. Eyes enlarged, often so much so as to distend
the head dorsally and laterally.
2. Modification of the rostrum, either thinner or
shorter, occasionally more elongate.
3. Fuzz or pubescence, often composed clearly of
aesthetascs, occurs medially on article 1 of antenna
1 and dorsomedially on articles 3-4 of antenna 2.
4. The ventral penicillate setules on article 1 of
antenna 1 greatly increase in number.
NUMBER 245 27
5. Aesthetascs occasionally become enlarged.
6. Calceoli develop on the primary flagellum of
antenna 1 and on the flagellum of antenna 2 and
often dorsally on article 5 of the peduncle on
antenna 2. Usually one calceolus occurs on each of
the first few articles of the primary flagellum of
antenna 1 and upwards to 7 occur dorsally on
article 5 of antenna 2. Clusters of male setae ac-
company the calceoli of article 5, often more clus-
ters than calceoli. Calceoli of antenna 2 are stated
for at least one male of each species where known
but these appear to be nondiagnostic at species
level in the form we quote because right and left
antennae may be in different phases of develop-
ment; apparently male flagellar articles proliferate
from article 1 and alternate in presence and ab-
sence of a calceolus so that the formulas are vari-
able within a species. Calceoli occur alternately to
the penultimate article of the flagellum in fully
terminal males but are rudimentary towards the
apex in younger males.
7. The flagellar articles of antenna 2 proliferate
and increase in length, thereby increasing the over-
all length of antenna 2, often exceeding the body
length.
8. Occasionally the upper lip becomes heavily
cornified with the ventral margin becoming either
less or more excavate than in the female.
9. Mandibular palp article 3 occasionally bears
supernumerary setae on the inner margin, and the
basofacial setae may become altered in number or
length.
10. Minor changes in other mouthparts infre-
quently occur, such as modification of the right
lacinia mobilis and parts of maxilla 1 and the
maxilliped.
11. Minor modifications in gnathopods and
pereopods 1-2 often occur, such as reduction in
numbers and lengths of setae.
12. Pereopods 3-5 are often modified to a small
degree, article 2 of pereopod 5 usually becoming
much narrower than in the female, less frequently
so on pereopods 3?4; article 6 of pereopod 3, and
especially of pereopod 4, becoming more elongate
than in the female but other articles rarely modi-
fied in proportion.
13. Males develop one penial process on each
side of the seventh thoracic sternite just medial to
the coxal articulation.
14. The pleopods become very powerful, with
stout peduncles and enlarged rami bearing highly
elongate setae; the peduncle of pleopod 3 in males
usually is significantly shorter than that of pleopods
1-2.
15. While the pleosome becomes more powerful
in male phoxocephalids the urosome usually dimin-
ishes in size, becomes thinner dorsoventrally and
often more elongate. Uropods 1-2 diminish in in
size concordantly.
16. Epimera 1-3 are usually broadened antero-
posteriorly and epimeron 3 occasionally is shortened
posteriorly. The posterior margins of epimera 1-2
often are grossly protuberant and smoother than in
the female, with setae on epimera 1-3 often de-
creasing in length and number.
17. Uropods 1-2 usually undergo changes in the
number and lengths of setae and spines, the most
conspicuous modification being the proliferation
and increase in length of spines on the peduncle
of uropod 2. The spines on the inner ramus of
uropod 2 are occasionally reduced or lost com-
pletely.
18. Uropod 3 of female phoxocephalids outside
of Australia usually has the inner ramus quite
short and poorly setose. In most Australian species
the female inner ramus is elongate more or less
as in the male, but males usually have the inner
ramus even more elongate, well exceeding article
1 of the outer ramus and usually far more setose;
the outer ramus is either more setose or actually
bears setae as well as spines (noting that some
females also have setae).
19. The telson is usually more elongate and occa-
sionally broader than in females, the apical spines
are shortened and a long irregular row or linear
patch of denticles develops dorsally and basally
near the midline of each lobe.
MINOR ATTRIBUTES.?The following characteris-
tics are relegated to a low level of importance.
Though most of them are illustrated for many
species they are often consistent from species to
species in a genus and unless yariability occurs
they are omitted from discussion in the sections
"Observations" and "Illustrations" when they have
not been illustrated for a particular taxon. Because
most cases of illustrative and descriptive omission
occur in the diverse genus Birubius the following
list of characters and comments primarily concerns
that genus.
Size or shape of cephalic protrusions adjacent to
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
eye and depth or shape of anteroventral corner of
head are mostly illustrated but, because of head
deformation after preservation and difficulty in
replication of precise angles of observation, these
characters have low taxonomic value. Minute mar-
ginal setae on the dorsal view of the rostrum are
omitted. Medial views of antennae are omitted
except in the case of an occasional article 1 of
antenna 1 of a male to show the medial patch of
fuzz. Article 1 of antenna 1 in the female bears a
midapical facial (lateral) setule; midapical setules
of articles 2-3 are few in number (3-4 each). The
distalmost seta on the lateral face of article 3 on
antenna 2 is usually much longer than the prox-
imal seta. Proximal spine groups on the lateral face
of article 4 on antenna 2 are accompanied by an
accessory setule that is always illustrated but not
counted in the spine formulas and otherwise ac-
corded no great taxonomic value. Ventral setal
groups usually have the subbasal pair of plusetules
omitted or reduced in prominence in the illustra-
tions, or are often hidden medially. Foreshortening
of facial spines is ignored. The apex on the
flagellum of male antenna 2 bears one elongate
seta and a short setule. The gland cone is hidden
between the juncture of articles 2 and 3. The
prebuccal complex is usually illustrated but de-
scribed if not illustrated: the lateral view is gener-
ally uniform in Australian species, any exceptions
are noted, the anterior view is variable from species
to species (see p. 13). The full body of the man-
dible with attached palp is omitted, the palpar
hump either being illustrated or described. Setules
between mandibular raker spines are deemphasized
and the texture of mandibular molars (extremely
weak in any event), is ignored. Numerous views of
the right lacinia mobilis (usually drawn from an
aspect attained when the raker spines are fully
extended), have been eliminated. Poor preservation
and fleshiness of lower lips reduces their value in
distinguishing taxa except for the presence or
absence of meatal cones. Except where mentioned
the outer plate of maxilla 1 is furnished with 11
spines in 2 groups of 6 and 5, the smaller group
with outer bifid apices, and spine 2 of 6-group
(second from lateral margin) thickened. Departures
from normal include reduction to 9 or 7 spines.
Right and left palps of maxilla 1 are approxi-
mately similar. Fuzz striations or seriate ranks on
maxillipedal palp article 4 are ommitted. "Or-
dinary" gnathopods denote gnathopod 1 slightly
longer and thinner in apical parts than gnathopod
2, and radical departure is so described, often in
the main body of the description. The precise shape
of crenulations on the palms of gnathopods has
not been depicted. Coxae are taller than broad,
coxae 1-4 with a small anteroventral bulbar setule,
coxa 4 with additional small posteroventral bulbar
setule. Departures occur mainly in those few species
with the setule not clearly bulbar or in the pres-
ence of supernumerary setules on coxa 4. Article 5
of pereopods 1-2 is furnished with a short spine
contiguous to the main apical spine. Pereopod 2
is usually not illustrated but in the main descrip-
tion setal formulas on articles 4-5 of pereopods
1-2 are quoted. Except where severaly altered,
dactyls of pereopods 3-5 have been ignored, al-
though they have setules and protrusions similar
to those found on pereopods 1-2 but in a degree
of miniaturization. The ventral and posterior
margins of article 2 on pereopods 5-5 are termed
naked unless some of the setules have become
elongate setae. Pleopods have been omitted en-
tirely.
Our assessment of certain characters may have
been defective. Plumes and disjunct spines on
mandibular molars or cuticular setules and orna-
ments are so small as to be at the limits of light
microscopy. Ventral setules on article 1 of antenna
1 are often damaged and subject to interpretation
as to how far ventrally a setule must be placed to
be included. Denticles on male telsons have not
been studied with oil-immersion microscopy as the
denticles are so small that superior results probably
will be obtained with scanning electron micro-
scopy.
Long descriptive essays on spine and setal lengths
and subtleties in shapes and sizes of other parts
are generally omitted, although these subtleties are
occasionally invoked to make comparisons between
species. Setal plumosities are omitted in the vast
majority of the illustrations. Discernible by re-
fracted light in microscopic views they are too fine
for accurate rendition. The presence or absence of
plumes on certain setal groups on various appen-
dages may someday be useful in studies of cryptic
or sibling speciation but this aspect has not been
pursued except in the Tipimeginae. Considerable
success in detecting distinct but closely similar
species has been possible with other characters.
NUMBER 245 29
An understanding of "microspeciation" must come
from studies utilizing vast materials from wide
geographic ranges.
SENILITY.?Amphipods apparently are subject to
the exigencies of senility, surviving beyond useful
reproductive stages and undergoing terminal dis-
placements of morphology, perhaps in a final ecdy-
sis. Several specimens in the collections supply weak
evidence that these terminal anomalies may occur
in reproductive adults as well as postparturative
individuals.
Extremely large adults are called "superfemales"
or "superadults" and those with aberrancies, espe-
cially the loss of brood plates, are termed "seniles."
Superadults are most easily recognized because of
gigantism; they may be 20-50 percent greater in
body length than ordinary adults and, of course,
are much more massive. In the literature they have
often been called "intersexes," but that term is re-
stricted herein to those individuals bearing both
penes and brood plates. No special search for inter-
sexes has been conducted in this study although 2
specimens of Birubius with both sexual character-
istics have been found.
Superadults often provide taxonomic difficulties
because of character displacement or character con-
vergence towards the typologically separated ordi-
nary individuals of similar species. Occasional mu-
seum collections are dominated by such individuals
in certain species because they are large, easily seen
with the naked eye and have been handpicked from
massive samples while the less conspicuous adults
of ordinary size have been discarded. Massive sam-
ples screened through wide meshes often contain
mostly superadults of amphipods. Because of fine-
ness of screening, the survey samples of Western
Port and Port Phillip Bay contain only a few super-
adults in proportion to normal adults thereby indi-
cating that they are an insignificant part of the
populations of phoxocephalids in those bays. These
superadults have usually been avoided herein as
typological models for taxonomic exposition but
frequently the complications of their morphological
anomalies to the orderly identification of species
are mentioned. For this reason they appear to occur
more prominently in this study than they actually
do in nature.
SUMMARY OF TAXONOMIC EVALUATIONS.?The
evolutionary trend within the Phoxocephalidae
matches that of other groups of amphipods and
that of the order as a whole. Hypothetically, phoxo-
cephalids commence in primitive state as fully de-
veloped fossorial organisms of large size with stout
body, heavily spinose and thickened antennae and
pereopods, and heavily spinose uropods. Then they
undergo the following modifications in various evo-
lutionary lines: reduction in fossorial mechanisms
through decrease in body size, streamlining of
pereopods, and reduction in spinosity. Primitive
gnathopods are a fully diverse pair with gnathopod
2 enlarged and its article 5 shortened, but the most
advanced members of most subfamilies have gnath-
opod 2 reduced in size and beginning to assume the
appearance of gnathopod 1. In some groups pygid-
ization is occurring, to wit, the loss of complexity
and articulation in the urosome through fusion of
urosomites or the inner ramus of uropod 2 to the
peduncle. One genus, Joubinella, has become fully
nektonic and other joubinellins may be in the proc-
ess of adapting to the inquilinous or commensalistic
mode in benthic habitats.
Phoxocephalid Environment
PHOXOCEPHALIDAE IN AUSTRALIA.?Phoxocepha-
lidae are a dominantly fossorial group of Amphi-
poda with limited neritic phases. Both females and
immature males are mainly benthic burrowers on
coastal shelf sediments that have size ranges in
median diameters of 0.12 to 0.93 pjn. Phoxocepha-
lids appear to be highly sensitive to any conditions
outside of the open sea and therefore are rarely
reported from embayments regardless of salinities.
Their primary evolutionary habitat has been on
the fine grained sediments of fully marine coastal
shelves (distributional data, J. L. Barnard, 1960).
Secondarily, Phoxocephalidae have been found
on intertidal sediments (for example, Microphoxus
minimus in Panama, J. L. Barnard, 1960) and occa-
sionally occur in intertidal samples of dominant
epiflora where they probably burrow into tiny
patches of trapped sediment (J. L. Barnard, 1969b).
The discovery of 16 genera and 54 species of
phoxocephalids in two embayments of Victoria is
of considerable interest in light of previous knowl-
edge about phoxocephalid ecology. In the sparse
records (Reish and Barnard, 1967, Barnard and
Reish, 1959, J. L. Barnard, 1964a, Bousfield, 1973)
no such similar embayment has ever been reported
to contain more than two or three species of phoxo-
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
cephalids. No other part of the world has such an
abundance of phoxocephalids in the shallow open
sea as does Australia, the next highest number being
the 21 species of California.
No unusual features have been discovered about
the environment of Port Phillip Bay and Western
Port to explain the high diversity of phoxocephalids
in those confined waters. By the standards known
for other world embayments one might describe
the Australian bays as "species packed." Not only
do these embayments contain many phoxocephalids
but they contain many other amphipods as well.
More than 150 species have already been identified
(only partly published as yet, see also J. L. Barnard,
1972a, 1974; M. M. Drummond in preparation).
This number approximates the total number of
shallow-water amphipod species in the entire warm-
temperate province of western North America and
far exceeds the 45 species found in Bahia de San
Quintfn, Mexico, by J. L. Barnard (1964a), an em-
bayment somewhat similar to Western Port.
The two bays of Victoria lie in the cool-temperate
(but not cold-temperate) Maugean province that in-
cludes Tasmania (Bennett and Pope, 1953, 1960,
Knox, 1963). Thermal levels taken by the WPBES
investigation in upper waters of adjacent Bass Strait
varied between 13.0? (winter) and 19.6?C (summer)
over the seasons. In Western Port water tempera-
tures varied between 10? and 21? but in shallower
waters they were 9.6? and 22.4? and on mudflats
7.8? and 29?. In Port Phillip Bay between 1947 and
1952 the means ranged between 7?-8? (winter) and
20? (summer) (Rochford, 1966). These values sug-
gest a cool but appropriately categorized warm-
temperate regime if the definition is limited to
thermal 10?-20?C for 9 months of the year. This is
the definition used in amphipod biogeography by
J. L. Barnard (1962, 1969a, 1972b).
Port Phillip Bay is a large, ovate bay of about
1950 km2, produced into a long western arm, and
with a narrow entrance 3.22 km wide. Its maximum
depth, occurring in a small area just inside the en-
trance, is about 91 m, but average depth is very
much less, and more than one-half of the total area
is less than 14 m deep. The bottom is generally
without major relief, sloping fairly evenly towards
a large central area with depths in the vicinity of
22 m.
Sediments are mainly clay and silty clay in the
central area, sand in the east and sooth*, with sedi-
ments intermediate between these in the northwest.
Macroscopic vegetation is fairly sparse except
around the periphery of the bay on submerged
rocky outcrops (PPBES Report on Phase 1, 1968-71,
Melbourne). Cell concentrations of phytoplankton
are not extraordinary. Water circulation in the bay
is slow, and total flushing takes five months. Salin-
ity levels are very close to those of the open sea.
Despite this wholly unextraordinary environ-
ment, there is sufficient environmental diversity to
enable at least 12 genera and 32 species of phoxo-
cephalids to occupy this small body of water.
Western Port is a much smaller, complex, tidal
embayment, about 1500 km3 in area, which includes
two large islands, the southernmost of which is so
placed that it divides the wide entrance into two
channels?the narrow Eastern, and the wider West-
ern, Entrance. The freshwater and sediment input
from the hinterland is very small. The water area
covers 680 km-, of which imertidal flats and sand
and mud banks account for 35 percent. The coast-
line, including the islands, is 263 km long. Sediment
is predominantly medium and fine sand, mixed
with varying proportions of clay (Marsden and
Mallett, 1975).
Compared with Port Phillip Bay, the bottom is
in much sharper relief; complex water movements
result in more varied sediment in the larger chan-
nels. Geologically, too, the areas are dissimilar
(Marsden and Mallett, 1975).
Dense beds of seagrasses cover about 38 percent
(260 km2) of the total area of the bottom of Western
Port. Zostera muelleri and Heterozostera tasmanica
extend over most of the intertidal flats; Amphibolous
antarctica is confined to a comparatively small area
in the main channel. Mangroves (Avicennia
marina), with salt marsh swamps, line 40 percent
of the 263 km of coastline and occupy an area of
12.1 km2. It is suggested (WPBES Report, 1975) that
the great extent of these two communities, seagrass
and mangrove, could significantly influence nutrient
cycling and productivity in the bay.
In light of the significant proportion of biotic
dominance by seagrasses, marshes, and mangroves,
and our knowledge of similar embayments such as
Morro Bay, California (Reish and Barnard, 1967),
Newport Bay, California (Barnard and Reish,
1959), and Bahia de San Quintin (J. L. Barnard,
1964a), the Amphipoda should be heavily domi-
nated (80 percent) by epifaunal rather than in-
NUMBER 245 31
faunal kinds. When contrasted with this expecta-
tion, the percent of infauna in the Australian embay-
ments appears strikingly high: we record 49 species
and 15 genera of phoxocephalids from Western
Port but have found only 70 species of 35 epifaunal
or nestling genera. At least 17 other species in 15
genera of infaunal or free living, nondomicolous
amphipods have also been collected in Western
Port.
Amphipoda generally appear to be more diverse
in southern Australia than in any other well studied
marine area (J. L. Barnard, 1969a). More than 300
species (including those presented herein) have been
reported from this area (Sheard, 1937, J. L. Bar-
nard, 1972a, 1974) and yet only a few families such
as the Dexaminidae and Phoxocephalidae have
been studied extensively in a modern context. At
least 50 family groups remain poorly studied and
will undoubtedly be found to contain hundreds
more of unknown species.
The high diversity of amphipods in southern
Australia suggests that the overall aspect of the
time-stability hypothesis (Sanders, 1969) is operable
in southern Australia but also that a factor of in-
cremental allopatric evolution must be integrated
into the system. The warm- and cool-temperate Aus-
tralian environment remains sufficiently steady over
long periods of time and is adequately diverse to
accumulate and preserve numerous species; but
those species must evolve and be naturally selected.
Such events producing new species require incre-
ments of instability sufficient to isolate populations
without damaging the general biotic stability neces-
sary to maintain the species already in existence.
Four kinds of isolation can be suggested. The
wide water distance between eastern Australia and
the predominantly warm-temperate fauna of New
Zealand is a possible divisory pathway. The enor-
mous distance between western and eastern Aus-
tralia separated by the environmentally undiverse
Australian Bight represents another isolating me-
chanism. If Lake Dieri flooded southern and mid-
dle Australia during Pliocene times, its runoff to
the sea could have further deteriorated an already
poorly diverse environment. The amphipod fauna of
southwestern Australia (J. L. Barnard, 1972a, 1974)
has many insular characteristics. It resembles the
fauna of New Zealand more than that of southeast-
ern Australia, thereby suggesting that it is func-
tionally isolated to some degree and may provide
a place, especially during elevated sealevels, where
division of pan-Australian populations may occur.
Finally, southeastern Australia has, or has had
in the past, numerous anchialine salt lakes such as
Lake King, almost fully disjunct from the sea but
providing environments in which amphipods, such
as Limnoporeia kingi, can survive and perhaps be
isolated for time sufficient to permit speciation.
Fluctuating sealevels would allow for isolation and
rejoining of these lakes to the sea.
BIOCEOCRAPHY OF PHOXOCEPHALIDAE (Figure 6).?
The compilation of genera and species in Figure 6
elucidates the dispersal of Phoxocephalidae from
an evolutionary center in Australia to other parts
of the world by disregarding early continental
aggregation. Data enclosed in boxes represent deep-
sea species for major ocean areas. Other data refer
to shallow-water phoxocephalids. The Magellanic
region and Falkland Islands represent one area,
the remainder of Antarctica, including the South
Georgia Islands (not included in Figure 6 but in-
dicated by an arrowhead on lower left margin),
represents a second area in the south. Other areas
are self evident. Paraphoxus oculatus is considered
to be a deep-water form.
Shallow-water Phoxocephalidae appear to be
more abundant and more diverse in Australia than
in any other part of the world. This broad state-
ment must be tempered strongly by the fact that
many areas of the world have either never been
explored adequately, or at all, for phoxocephalids.
Two areas, the northeastern Atlantic and southern
California, have been explored minutely but
phoxocephalids in the eastern Atlantic are about
one eighth as diverse as in Australia and in the
southern California area about one third as diverse
as in Australia.
The Okhotsk Sea has been studied extensively
with the conclusion that phoxocephalids are sparse
there. Only two species occur in South Africa,
another area now minutely explored. The numer-
ous publications on Indian and Ceylonese am-
phipods report only one species from those tropical
shores.
Antarctic and Magellanic regions have been
widely studied but the number of reported phoxo-
cephalids is fairly low; there is the possibility that
survey techniques there failed to recover soft
bottom amphipods in proportions equal to those
from other habitats. The Magellanic region ap-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 6.?World dispersal of Phoxocephalidae from Australian center (C = genera. S species.
E - endemic species (where pertinent to text), numerals indicate number of genera or species).
pears to be a major stopover on the long dispersal
journey from Australia to the Californias and
would be expected to have at least as many genera
and species as does California (see Figure 6).
Nevertheless, there is no evidence, other than in
Magellanic South America, that phoxocephalids
will be found elsewhere in terms of the vast abun-
dance now known in Australia.
Phoxocephalidae in the tropical shallows of the
Indo-Pacific region are sparse and apparently
dominated by the subfamily Phoxocephalinae as is
the case in Madagascar or by Mandibulophoxus in
India. The muddy east tropical Pacific shallows
have representatives mainly of Birubiinae such as
"Paraphoxus" floridanus or Microphoxus but the
Galapagos Islands have a shallow-water species of
Proharpinia, the northernmost oculate shallow-
water member of Harpiniinae except for Hetero-
phoxus oculatus which penetrates North American
waters as far as Oregon.
The California warm-temperate province and its
neighbors to the north and south support a rich
phoxocephalid fauna dominated by Birubiinae and
by species such as "Paraphoxus" fatigans and "P."
daboius (which probably should be removed to a
new subfamily once the several new genera con-
taining these species are described). None is a
member of Birubius. Many of the eastern Pacific
species in the undescribed subfamily bear weakly
ensiform antenna 2 and one group carries tri-
chophoxin gnathopods and has weakly enlarged
article 3 on pereopod 5 but otherwise is distinct
from trichophoxins. These groups are furnished
with the elongate article 2 of antenna 1 suggesting
relationship to birubiins rather than to brolgins
but several species have reduced spination on the
mandibular molars as in brolgins "Paraphoxus"
cognatus appears to belong to a new genus probably
allied to paraphoxins. California is therefore dom-
inated by Birubiinae and a second new subfamily
allied to Birubiinae, but, unlike Australia, it has
few shallow-water paraphoxins and only one
shallow-water harpiniin (Heterophoxus oculatus).
The Magellanic and Antarctic sectors appear to
have four brolgins (WildusT fuegiensis, Parharpinia
obliqua, P. rotundifrons, Pontharpinia uncinata
[the last three of indeterminant generic placement]);
two birubiins (Metharpinia longirostris and M.T
cornuta [probably distinct generically]); one par-
harpiniin {Parharpinia sinuata); two species of
Heterophoxus; two species of Proharpinia; and
one species of Coxophoxus. The influence of
brolgins from Australia is evident and the only
parharpiniin to be recorded outside of Australia
NUMBER 245
occurs in the Magellanic sector. Shallow-water
harpiniins are also a predominant group here,
the only place outside of the Galapagos Islands,
New Zealand, northern Atlantic, or northwestern
America where this group lives on coastal shelves.
New Zealand probably has a rich phoxocephalid
element. Nine species in eight genera, (of which
only five are established) have already been de-
scribed. The fauna of New Zealand is strongly
related to that of Australia as shown by the occur-
rence of Trichophoxus, a sibling genus of Tipi-
megus; Protophoxus, a partner of Parharpinia; two
genera represented by Waitangi rakiura, Para-
phoxus chelatus, and P. spinibasus, again repre-
senting trichophoxins; a brolgin, U'ildus waipiro;
Paraphoxus pyripes, which appears to belong to a
new genus of unknown affinity perhaps related to
parharpiniins or brolgins; and one species each of
Phoxocephalus and Proharpinia. The latter har-
piniin reflects the subantarctic, non-Australian ele-
ment in New Zealand.
The northwestern Pacific shallows, including
Japan and the Okhotsk Sea, have eight species
primarily of birubiins in several new genera, plus
one brolgin. The relationship to the northeastern
Pacific is strong. The origins of this group, either
across tropical shallows from Australia or against
the prevailing southeastward oceanic drift from
America, are obscure.
Western Atlantic shallows are poorly explored
except in the.New England area. The cold north
has the predominantly north Atlantic Phoxo-
cephalus holbolli and, apart from seven species of
Harpinia, the other two known species ("Para-
phoxus" epistomus and "P." spinosus) have siblings
found in the eastern Pacific Ocean, suggesting that
they passed through the mid-American isthmus in
an eastward and northward direction.
The northeastern Atlantic Ocean and Mediter-
ranean Sea are well studied but generically impov-
erished. Only Metaphoxus, Phoxocephalus, and
Harpinia live in those seas. Harpinia is the domi-
nant genus and is probably endemic to the area or,
at most, has escaped by a polar route to high lati-
tudes of the north Pacific. Harpinia is one of the
most advanced and specialized genera of phoxo-
cephalids; apparently its origins lie in the sub-
antarctic harpiniin stock and its migration occurred
via a bathyal pathway (as indicated by its blind-
ness).
The deep seas are dominated by blind harpiniins
and a few other genera mainly of the subfamily
Phoxocephalinae. Joubinella is found in bathy-
pelagic waters but net tows through upper bathy-
pelagic and neritic waters often capture nektonic
males of various brolgin and phoxocephalin genera.
In summary, the great diversity of phoxocepha-
lids in Australia, the limited generic diversity and
great morphological advancement of phoxocepha-
lids in the northeastern Atlantic, and the domi-
nance of the deep sea by harpiniins whose shallow-
water representatives today live mainly in Sub-
antarctica, suggest that Phoxocephalidae underwent
their major subfamilial elaboration in the southern
hemisphere and are in the process of invading fav-
orable environments of the northern hemisphere.
They may be meeting stiff competition from other
fossorial families such as urothoids, pontoporeiids,
and oedicerotids and may have withdrawn to some
extent against an outward flow of those groups; but
this appears unlikely in view of the fact that a
subantarctic group of blind harpiniins has popu-
lated deep sublittoral waters of the northeastern
Atlantic. This odd occurrence suggests that the
pathway for this immigration may have been the
deep sea. Few oculate phoxocephalids have managed
to overcome whatever barriers exist to keep them
out of the northeastern Atlantic via a shallow-water
route from the south. One of those has defective
eyes {Phoxocephalus holbolli) and therefore may
have moved along a bathyal route.
One assumes that the Magellanic fauna is a de-
scendant of the Australian fauna through the West
Wind Drift or from original Gondwana connec-
tions. From South America, phoxocephalids could
have moved into the Antarctic and northward
through the constricted east Pacific tropics charac-
terized by muddy shores and upwelling.
The west coast of the Americas therefore appears
to have been a favorable pathway for phoxocepha-
lids to move northward and to populate the Cali-
fornias richly. Some of those species have close rela-
tionships with the Japan-Okhotsk sector and may
have immigrated from that region; but the opposite
migratory path is more plausible than a path from
Australia through east Asian seas. Several species
passed into the western Atlantic during the opening
of isthmian waterways.
Phoxocephalids have not succeeded in the tropics
except in Madagascar and in the muds of the east-
34
ern Pacific. South Africa has been strongly isolated
from phoxocephalids. The west coast of Africa and
lower latitudes of South America are poorly known.
In so many distribution patterns of this kind
where fossil evidence is lacking one may hypothesize
two striking alternatives: (1) the Phoxocephalidae
are now being constrained into a relict position and
are surviving best in the deep sea, in the otherwise
impoverished areas (for amphipods) such as the
northeastern Pacific, or in highly isolated regions
such as warm-temperate Australia and cold-
temperate South America; or (2) the Phoxocephali-
dae are a relatively youthful group blossoming in
the southern hemisphere, and are in the process of
immigrating into the northern hemisphere and in-
vading the relatively youthful and very cold deep
seas. Morphologically, phoxocephalids appear to be
very advanced and efficient organisms, of late origin.
Though fossorial, they, unlike any ha us tori ids or
few pontoporeiids and urothoids, have both invaded
the deep sea and developed numerous species of
reduced fossorial appearance which may be adapted
to a semifossorial role in their habitats. Not only
the severe reduction of rostra but especially the
development of non-skid cuticle in several of the
Australian taxa suggest that the family is beginning
to move out of its primarily fossorial role into an
ecological position broadly known as inquilinous.
Phoxocephalid Evolution
ORIGIN OF PHOXOCEPHALIDAE.?Phoxocephalidae
appear to have their closest direct morphological
resemblance to Ponto-Caspian genera of the Gam-
maridae (sensu lato) and somewhat more remote
resemblance to the haustoriid-urothoid-pontoporeiid
group of families.
A cluster of genera to be called Pontogammarids
and comprising genera such as Niphargoides Sars,
Niphargogammarus Birstein, Uroniphargoides
Stock, Compactogammarus Stock, Obesogammarus
Stock, Paraniphargoides Stock, and Pontogammarus
Sowinsky represent the focus of this discussion.
Other genera belong to this group but lie outside
the needs of the discussion. The group has been
revised taxonomically by Stock (1974).
The Pontogammarids appear to contain the fol-
lowing attributes suggesting an evolutionary de-
scent towards phoxocephalids: (1) fossorial pereo-
pods 1-5; (2) shortened article 2 of antenna 1 with
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
apicoventral setation; (3) rich development of peni-
cillate setules ventrally on article 1 of antenna 1;
(4) elongate accessory flagellum; (5) reduction in
size of article 3 of antenna 2; (6) dominance of
antenna 2 by articles 4-5 of the peduncle but with
a weakly expressed phoxocephalid tendency for
reduction in article 5; (7) expansion of article 1
on antenna 2; (8) retention of the enlarged gna-
thopod 2 with shortness of article 5: (9) shape and
setation of pereopods 1-2; (10) expansion of
article 2 on pereopod 5 with facial setation, often
in the form of discrete setular bundles; (11) pres-
ence of facial setae on epimera; (12) continuous
dorsal spination or at least the presence of acces-
sory apical nails on the rami of uropods 1-2; (IS)
dominance of the outer ramus on uropod 3, with a
distinct second article and somewhat to greatly
shortened and subsidiary inner ramus, the rami
elongate and lanceolate; (14) deeply cleft telson;
(15) well developed inner lobes of the lower lip;
(16) reduction of setae on the inner plate of maxilla
1 (infrequent).
Not all of these are characteristic of all Ponto-
Caspian genera cited. In Niphargogammarus, for
example, a protophoxocephalid morphology is re-
flected in the shortened article 3 of antenna 2, the
well expanded article 2 of pereopod 5, and the
shape of antenna 1. In Compactogammarus, an-
tennae 1-2, mandibles (in the loss of C and D
setae, see Stock 1974), and pereopod 5 are significant
in this context, gnathopod 2 only weakly.
All Phoxocephalidae differ from Ponto-Caspian
gammaroids in the development of a rostrum,
though this has become reduced in highly derived
genera or species which can be linked hypotheti-
cally to fully rostrate ancestors. All phoxocephalids
have shortened articles 4-6 of pereopod 5 and
article 2 extends even more distally and posteriorly
than in Ponto-Caspian gammaroids. The phoxo-
cephalid mandibular palp lacks an expanded apex
on article 3. In no phoxocephalid is article 5 lobed
on gnathopod 2 whereas all pontogammarids have
a medium to strong lobe. Most phoxocephalids
carry gills on pereonite 7 but one subfamily, Tipi-
meginae, and the genus Limnoporeia, often lack
these gills. No phoxocephalid bears a submarginal
oblique row of setae on the inner plate of maxilla
1 but some of the more advanced pontogammarids
have also lost these setae. The inner plate of maxilla
1 in phoxocephalids always lacks medial setae and
NUMBER 245 35
rarely has more than 4 apical setae. All phoxo-
cephalids bear facial spines on article 4 of antenna
2, though in the more advanced members these
often become highly distal and scarcely facial.
Only one pontogammarid, a species of Niphargo-
gammarus, is known to have facial spines on arti-
cle 4 of antenna 2.
We hypothesize that a model ancestor of phoxo-
cephalids would bear a fully triturative molar and
enlarged gnathopod 2, similar to primitive gam-
maroids, many pontogammarids, and to primitive
living phoxocephalids, which we have identified as
Pontharpinia. The enlarged gnathopod 2, however,
is rapidly reduced in most phoxocephalid lines of
evolution and even in several pontogammarids.
Only five genera of phoxocephalids have a fully
triturative molar: Pontharpinia, Phoxocephalus,
Leptophoxoides, Joubinella, and Jerilderia and
each of these also has an enlarged gnathopod 2;
but in all of them article 5 is especially short, un-
lobed and essentially cryptic, unlike that of any
pontogammarid. Neither Phoxocephalus nor Jeril-
deria can be considered close to the ancestral type
because of the loss of segmentation in the palp of
maxilla 1. Joubinella is a highly adapted nektonic
genus, and Leptophoxoides is an apomorphic
abyssal genus with heavily modified maxilla 2 and
maxilliped.
Pontharpinia comes closer to the ancestral grade
than do the other extant genera because of the
biarticulate palp on maxilla 1, the supernumerary
setae on the inner plate of maxilla 1, the fully
setose maxilla 2 (but lacking oblique facial row),
the fully developed maxilliped, the rich setosity of
the mandibular palp (but C and D setae absent),
the presence of more than 2 setae or spines on
article 3 of antenna 2, the absence of cones on the
lower lip, the presence of facial setae in bundles on
pereopod 5, the somewhat elongate articles 4-6 of
pereopod 5, the presence of setosity on article 2 of
pereopods 3-4, the presence of facial setae on
epimeron 3, the apically spinose rami of uropods
1?2, and like many other primitive Australian
phoxocephalids but not necessarily like ponto-
gammarids, the fully magniramus uropod 3 with
elongate and setose inner ramus in the female. The
telson of Pontharpinia is heavily specialized and
divergent from pontogammarids and other phoxo-
cephalids.
As has been noted above, in those genera of
phoxocephalids with enlarged gnathopod 2, article
5 is nonlobate, and as in Pontharpinia, is usually
cryptic, unlike any pontogammarids. If phoxo-
cephalids with enlarged gnathopod 2 are considered
to be those genera with strongest ancestral links,
then the more advanced phoxocephalids generally
have undergone a reversion to the elongate con-
dition of article 5 concomitant with reduction in
size of gnathopod 2. This condition is similar to
many gammaroids and those few pontogammarids
with reduced gnathopod 2.
Phoxocephalids have many characters in com-
mon with one or more genera of haustoriids, ponto-
poreiids, and urothoids (the term "haustorioid" is
used hereafter for this group of families). All
haustorioids are characterized by reduced, almost
mitten-shaped gnathopods; none bears the enlarged
gnathopod 2 seen in various primitive phoxo-
cephalids. Most phoxocephalids bear somewhat
broadened hands with well developed dactyls fitting
the palms, except for the Tipimeginae and a new
genus typified by "Paraphoxus" variatus. Few haus-
torioids (except Zobracho) have a rostrum as well
developed as that found in Phoxocephalidae.
Haustorioids and phoxocephalids share similar
fossorial pereopods, antennae, and uropods but no
haustorioid is admitted to the Phoxocephalidae
because of the specific combination of the following
characters in Phoxocephalidae: visor-like rostrum
with expanded, deltoid article 2 of pereopod 5,
and shortened stenopodous articles 4-6. No haus-
torioid bears precisely the kind of triturative man-
dibular molar found in primitive phoxocephalids
(no pontogammarid bears this kind either), whereas
a few haustorioids bear the enlarged kind found in
pontogammarids or have an immensely enlarged
and fuzzy molar. Many haustorioids have variously
modified coxae whereas the coxae of Phoxocepha-
lidae fit a consistent, almost invariable basic pat-
tern. Many haustorioids have fully setose inner
plates of maxilla 1 and have the oblique facial
row of setae on the inner plate of maxilla 2. A few
haustorioids carry facial spines on article 4 of
antenna 2. Several have poorly cleft or greatly
shortened telsons and pontogammarid-like man-
dibular palps or shortened accessory flagella.
Although phoxocephalids, pontoporeiids, haus-
toriids, and urothoids may have a common ancestry
and this ancestral line may have proceeded out-
ward from a gammarid precursor similar to the
36
pontogammarid form, the direct linkages between
phoxocephalids and haustorioids (sensu lato) have
not been preserved in the modern fauna (as known
at this time). Phoxocephalids generally retain
plesiomorphic gnathopods, whereas haustorioids
retain plesiomorphic maxillae on which numerous
specializations have been superimposed. Despite the
vast degree of convergence among the various
groups, almost every genus of both phoxocephalids
and haustorioids can be compared minutely and
found to have characters which prevent a simple,
parsimonious, incremental evolutionary system to
be erected between the groups. For example, Uro-
thoe is furnished with a rostral area and pereopod
5 of incipient phoxocephalid character but the
mandibular incisors are highly apomorphic, where-
as phoxocephalids invariably possess plesiomorphic
incisors.
A model ancestor can be hypothesized to diverge
into both phoxocephalids and urothoids, or into
pontoporeiids and phoxocephalids; but this ances-
tor remains as close morphologically to ponto-
gammarids as it does to pontoporeiids, urothoids,
or phoxocephalids. The conclusion is reached that
erection of a model ancestor for phoxocephalids
within those groups, or extrapolation from Phoxo-
cephalidae of an ancestor for haustorioids is fruit-
less in light of present knowledge. Ancestry in
organisms like pontogammarids for at least ponto-
poreiids and phoxocephalids remains the most
parsimonious solution in light of known mor-
phology.
The relationship of Phoxocephalidae to Ponto-
Caspian gammarids is tenuous geographically. The
nearby Mediterranean Sea and eastern Atlantic
Ocean are greatly impoverished of Phoxocephalidae;
apparently all eastern Atlantic phoxocephalids are
highly advanced members of the family. If phoxo-
cephalids did descend from early pontogammarids
the thesis must be proposed that a pontogammarid
escaped eastward through a Tethyan seaway to an
Australian-like environment highly remote from
the Ponto-Caspian basin and Mediterranean Sea.
Many increments of evolution then took place, but
the ensuing barrier between the Ponto-Caspian
Basin and the phoxocephalid center has prevented
more than a token return of phoxcephalids into
the adjoining modern sea basins. This barrier was
probably a complex of thermal, competitive, and
geographic elements. Perhaps drying of the Medi-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
terranean basin in the Pliocene extinguished any
surviving links between pontogammarids and
phoxocephalids, although other groups such as
sarothrogammarids survived in the trans-Gibraltar
eastern Atlantic (J. L. Barnard, in prep.).
The modern position of continents and thermal
regimes stretches credibility that phoxocephalids,
which appear primarily adapted to nontropical
environments, could pass easily from the south
warm-temperate through the tropical regime to
northern cool waters. One might therefore suppose
that the protophoxocephalids would have had an
equally difficult time passing from Ponto-Caspian
regions into the southern hemisphere, and there-
fore one might suggest that the events required
positions of continents and mild (not torrid)
thermal regimes conducive to a continuous environ-
mental pathway from the Ponto-Caspian basins to
the Australian regime.
Modern environments do not appear favorable
to widespread dispersal of phoxocephalids away
from Australia, suggesting that the attenuated
faunas of many areas are the result of a coincidence
between the major events in phoxocephalid evolu-
tion in Australia and environmental conditions
more or less similar to those now prevailing.
Phoxocephalids are burrowers. Shallow-water
species might be expected to have more difficulties
in dispersing by rafted materials than nestling or
domicolous amphipods. Although the West Wind
Drift is a good transporting mechanism and may
have brought about a connection between Aus-
tralia and South America, the physical difficulties
encountered by a fossorial amphipod in such a
migration must be enormous. Dispersal of shallow-
water phoxocephalids is probably hampered by
long sea distances and torrid tropical waters which
suggest that Australia moved into its present posi-
tion and marine environmental regime after the
protophoxocephalid reached those shores but be-
fore much outward dispersal of the group had oc-
curred. The poor return of phoxocephalids from
Australia to the northern hemisphere suggests that
the Australia-South America connection through
Africa in the time of Gondwanaland persisted long
after the arrival of phoxocephalids.
The very impoverished situation in South Africa
today and the sparse fauna of South America, how-
ever, suggest either that Australia was already iso-
lated before the phoxocephalid arrival and that
NUMBER 245 37
modern distributions reflect dispersal across wide
water gaps, or that once split away from Australia,
South African and South American sublittoral
zones deteriorated environmentally.
EVOLUTIONARY DESCENT IN PHOXOCEPHALIDAE
(Figure 7).?If Pontharpinia is taken as a model
approaching the primitive phoxocephalid, the lines
of descent to various more advanced subfamilies
are marked by the following trends.
Tipimeginae: Reduction in overall size of
pereopod 5, with increase in size of article 3, loss
of posteroproximal setae on article 5 of pereopods
1-2, elongation of article 2 on antenna 1, loss of
trituration on molar, resulting in a pseudotritura-
tive form, reduction in size of both gnathopods,
almost mitten-shaped with thin hands bearing
heavy anterior setation, and development on uro-
pod 1 of a large ventral spike. The Tipimeginae
retain facial setae on pereopod 5, dispersed facial
setae on palp article 3 of the maxilliped, numerous
ornaments on article 3 of antenna 2, extra apical
setation on article 2 of the outer ramus on uropod
3, facial and posterior setae on the epimera, broadly
fossorial pereopods 3-4 and a relatively enlarged
molar bearing 3-4 enlarged spines and having the
appearance of being derived from the cusps of a
triturative molar. Tipimegins also have an enlarged
midapical spine on article 6 of pereopods 1-2,and a
medial comb on article 5 of those pereopods. An-
tenna 2 is weakly ensiform and coxa 4 is heavily
setose posteriorly.
Brolginae: Reduction of mandibular molar to a
small hump bearing 3 or fewer elongate spines
more or less congealed basally, often with a basal
plate, reduction of gnathopod 2 in advanced mem-
bers with elongation of article 5, reduction in
breadth of articles 4-5 on pereopods 4-5, simplifica-
tion of epimeron 3, with loss of all but a few setae
or setules, smooth rounding of the posterior margin,
and occasional retention of a special peduncular
spine on uropod 1.
Leongathinae: Reduction in mandibular molar
to a semitriturative condition in which one edge of
the discoid molar is toothed much as one side of a
triturative molar is toothed.
Joubinellinae: Weak to strong elongation of
article 2 on antenna 1, reduction of molar in ad-
vanced members to a nontriturative hump with
articulate spines.
Parharpiniinae: Reduction of mandibular molar
and gnathopod 2, elongation of article 5 on gnatho-
pod 2, development of supernumerary dorsal spina-
tion on telson, retention of special spine on pedun-
cle of uropod 1, and distal tapering of article 2 on
pereopod 3.
Birubiinae: Weak to strong elongation of arti-
cle 2 on antenna 1, reduction of mandibular molar
to a nontriturative hump with articulate spines,
reduction in size of gnathopod 2 with elongation of
article 5.
Phoxocephalinae: Loss of segmentation in palp
of maxilla 1, simplification of epimeron 3 and re-
duction in setation on maxilla 2.
Harpiniinae: Reduction in width of article 2 on
pereopod 3, and, in advanced members, the develop-
ment of an ensiform process on antenna 2.
The general trends in phoxocephalid evolution
therefore appear to be reduction in complexities of
setation or spination on most parts of the body
thought to have fossorial importance, such as the
antennae, pereopods 1-5, uropods 1-2, mandibu-
lar palp and epimera, the reduction of molarial
complexities, loss of palpar segmentation on
maxilla 1, loss of setae on inner plate of maxilla 1
and the face of maxillipedal palp article 3, on the
mandible and maxilla 2, reduction in numbers of
spines on the outer plate of maxilla 1, development
of salivary cusps (cones) on the lower lip, reduction
in thickness of mandibular palp, reduction in size
of gnathopod 2 with reversional elongation of arti-
cle 5, loss of thickness on articles 4-5 of pereopods
3-4 and on article 2 of pereopod 3, loss of the spe-
cial spine on the peduncle of uropod 1, and reduc-
tion in body size.
Within certain evolutionary clusters such as Biru-
biinae a trend occurs towards the reduction of the
rostrum and loss of segmentation on the urosome.
In the Phoxocephalinae there appears to be a trend
towards elongation of the telson. In the Harpini-
inae a significant number of species has developed
macrocuspidation on article 2 of pereopod 5 and
the ensiform process on antenna 2 reaches its maxi-
mum development and is retained even in the most
advanced taxa such as Harpinia.
The deep-sea benthos has been invaded almost ex-
clusively by the Phoxocephalinae and Harpiniinae.
Genera of the latter subfamily are found in the
deep sea or in cold north Atlantic middepths. The
loss of eyes apparently is universal in species in-
habiting depths in excess of 600 m; body size is re-
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
? = ?
PONTHARPINIINAE
BIRUBIINAE
20 Birubius
21 Yan
26 Microphoxus
22 Tickalerus
1
23 Kulgaphoxus borralus
24 Kulgaphoxus cadgeeus
26 Metharpinia
s Mandibulophoxut
PHOXOCEPHALINAE
27 Phoxocephalu* I
28 Jerilderia
29 Leptophoxoidei
so Leptophoxut
VI
r
36 Japara
31 Metaphoxua
37 Rikkarus J/
38 Limnoporeia
\
39 Kondoleus
Metaphoxoidet
Diogodia*
Va$co
36 Hopiphoxus
44 Heterophoxus
?
46 Pseudharpinia 46 Harpiniopgis
47 Harpinia
FIGURE 7.?Evolutionary pattern of subfamilies and genera (except Waitangi) of Phoxocephalidae
(z ? unreduced; short thick arrow points to long or short article 3 of pereopod 5; for identifica-
tion of other symbol labels in key see "Figure Abbreviations").
NUMBER 845 39
duced and almost all the supposed fossorial mecha-
nisms are weakened, apparently as an adaptation to
fineness of sediments.
In one of the new Californian genera typified by
"Paraphoxus" bicuspidatus epimeron 3 becomes
shortened posteriorly in females as well as males;
this trend occurs in Australian males of Birubius.
Australian phoxocephalids exhibit all of the evo-
lutionary trends seen in exotic genera except for
the full development of an ensiform process on an-
tenna 2 and enlarged cusps of pereopod 5. These
latter developments are confined to subantarctic
shallow water Harpiniinae and their deep-sea de-
scendants.
PHOXOCEPHALIDAE
DIAGNOSIS.?Head elongate or shortened (special-
ized) but flat dorsoventrally and usually bearing a
flat, sagi tally arched, visor-like rostrum, this occa-
sionally obsolescent, lacking ventral retrorse pro-
trusion. Eyes when present ommatidial. Article 1
of antenna I elongate, accessory flagellum multi-
articulate. Mandibular molar small, either fully
triturative or nontriturative and bearing spines,
palp present, 3-articulate. Lower lip with strong in-
ner and mandibular lobes. Palp of maxilla 1 well
developed. Maxilliped with 4-articulate palp. An-
terior coxae of normal length or coxae 1-3 never
significantly and individually reduced in size; basic
gill formula, 2-7 but occasionally 3-7 or 2-6.
Gnathopods either subchelate or chelate, wrists
poorly lobate or not lobate, gnathopods never mit-
ten shaped. Pereopods fossorial, heavily spinose
and/or setose; only pereopod 4 (rarely 3) elongate,
pereopod 5 much shorter than pereopod 4, article 2
enlarged, shield-like, articles 4-6 thin relative to
article 2; pereopod 5 strongly distinct in size and
shape from pereopod 4. Uropod 3 biramous. Telson
laminar, cleft.
DESCRIPTION.?Body stout to medium in thick-
ness, anterior pereonites not strongly compressed
forward; body never carinate or processiferous an-
terior to urosomite 3. Peduncle of antenna 2 with
facial spines (except Joubinella). Mouthpart field
quadratiform. Inner plate of maxilla 1 poorly
setose, usually with 4 or fewer apical setae (primi-
tive forms with 6 small setae also mostly apical).
Inner plate of maxilla 2 rarely with fully medial
setae, never with oblique submarginal row of setae.
Outer plate of maxilliped always developed and
spinose. Urosomites free or fused together in vary-
ing degree.
SEXUAL DIMORPHISM.?Major distinctions con-
fined to elongation of flagellum on male antenna 2,
increase in size of peduncle on antenna 2, develop-
ment of calceoli, development of male setular bun-
dles on antenna 2 (rarely on antenna 1), occasional
increase in size of aesthetascs or rarely change in
sizes of articles on antenna 1; increase in size of
eyes; uropod 3 of female usually with short, poorly
setose inner ramus, elongate in male, both rami
more setose in male; pleopods becoming more
powerful with peduncle of pleopod 3 somewhat
shortened; body somewhat smaller but more stream-
lined than in female; urosome reduced and pleo-
some increased in size relative to female. Minor
differences including increased setation on impor-
tant swimming appendages but reduction of setae
where streamline flow impeded (epimera for exam-
ple); gnathopodal distinctions virtually absent.
TVPE GENUS.?Phoxocephalus Stebbing.
COMPOSITION.?The following new subfamilies:
Birubiinae, Brolginae, Harpiniinae, Joubinellinae,
Leongathinae, Parharpiniinae, Phoxocephalinae,
Pontharpiniinae, Tipimeginae.
REMARKS.?In all but a few genera or species this
family is easily recognized by the large dorsally
compressed rostrum, the characteristic shape and
small size of pereopod 5 relative to pereopod 4 and
in the presence of facial spines on the peduncle of
antenna 2. One genus, Joubinella, lacks facial spines
on antenna 2 but retains a large rostrum, while sev-
eral genera and species have reduced rostrum but
retain the characteristic pereopods 4-5 and facial
spines on antenna 2. The combination of any two
of these characters serves to distinguish phoxocepha-
lids from similar genera in the gammaroid-hausto-
rioid-platyischnopid family clusters.
Key to the Subfamilies of Phoxocephalidae
1. Article 2 of pereopod 3 of thin form HARPINHNAE
Article 2 of pereopod 3 of broad form 2
2. Palp of maxilla 1 uniarticulate PHOXOCEPHALINAE
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Palp of maxilla 1 biarticulate 5
3. Article 3 of percopod 5 enlarged, mandibular molar pseudotriturative, of form C (Figures
5: c, 16) TIMMECINAE
Article 3 of pereopod 5 not enlarged, mandibular molar not pseudoiriturative 4
4. Mandibular molar semitriturative, of form B (Figures 5: B. 58) LEONGATHINAE
Mandibular molar not semitriturate 5
5. Article 2 of antenna 1 especially shortened 6
Article 2 of antenna 1 elongate 8
6. Mandibular molar triturative, of form A (Figures 5: A, 8) PONTHAWINIINA*
Mandibular molar not triturative 7
7. Mandibular molar of special form E (Figure* 5: E, 37) bearing 3 or fewer spinet usually
with common base, article 2 of pereopod 3 not tapering distally, telson lacking super-
numerary dorsal spination BROLCINAK
Mandibular molar of form D (Figures 5: D, 105) bearing articulate spines not all organized
into common base, usually 4 or more widely spread spines, article 2 pereopod 3 tapering
distally, telson bearing supernumerary dorsal spination PAHHAKPINIINAE
8. Gnathopods 1 or 2 or both enlarged JOUBINELLINAE
Gnathopods 1 and 2 both small BWUBIINAE
PONTHARPINIINAE
DIAGNOSIS.?Article 2 of antenna 1 shortened.
Mandibular molar fully triturative. Palp of maxilla
1 biarticulate. Setation on maxilla 2 ordinary. Gna-
thopod 2 significantly enlarged. Article 2 of pereo-
pod 3 of broad form; pereopod 5 with slightly en-
larged article 3.
SPECIAL CHARACTER.?Telson with large setal
brushes.
DESCRIPTION.?Article 5 of antenna 2 reduced.
Epimeron 3 of nonrounded classification. Apices of
peduncles on uropods 1-2 not combed.
TYPE GENUS.?Pontharpinia Stebbing.
COMPOSITION.?Unique.
Pontharpinia Stebbing
Pontharpinia Stebbing, 1897:32-33.
DIAGNOSIS.?Eyes present. Flagella of antennae 1-2
unreduced in female; article 2 of antenna 1 espe-
cially shortened, ventral setae widely spread; article
1 of antenna 2 weakly ensiform, article 3 with nu-
merous setae and setules, facial spines on article 4
in 2 or more rows plus special apical long spines
and short posterior spines, article 5 especially thin.
Right mandibular incisor with 3 teeth, molar
strongly triturative, medium, usually bearing fuzz,
palpar hump medium. Palp of maxilla I biarticu-
late, inner plate with 5+ setae. Setation of maxilla
2 ordinary. Inner plate of maxillipeds ordinary,
apex of palp article 3 weakly protuberant, dactyl
elongate, apical nail distinct, mostly immersed,
short. Gnathopods dissimilar; gnathopod 2 strongly
enlarged; article 5 of gnathopod 1 of ordinary
length, free, on gnathopod 2, cryptic, with weak
eusirid attachment, palms oblique, hand of gnatho-
pod 2 broadened; hands of both gnathopods poorly
setose anteriorly. Article 5 of pereopods 1-2 not
setose posteroproximally. Article 2 of pereopod S of
broad form; articles 4-5 of pereopods 3-4 broad to
medium, article 2 of pereopods 3-4 setose posteri-
orly; pereopod 5 ordinary, article 2 strongly setose
ventrally, article 3 slightly enlarged, dactyl normal
but with supernumerary setae. Epimera 1-2 lacking
long posterior setae, without midfacial setae above
ventral facial ridge except weakly on epimeron 2;
epimeron 3 ordinary. Peduncle of uropod 1 norm-
ally elongate, without apicoventral spike, with spe-
cial enlarged apicomedial spine, peduncular apices
of uropods 1-2 not combed, inner ramus of uropod
1 with 2 rows of marginal spines, all rami continu-
ously spinose to apex, inner ramus of uropod 2
ordinary. Uropod 3 ordinary, article 2 of outer
ramus carrying 3 short, apical setae. Telson with
only one apicolateral spine on each lobe plus
setules, with special brush of dorsal and lateral
setae. Urosomite 1 bearing one or more midventral
crescents or bundles of setae and sparse apicoven-
tral setae near base of uropod 1; urosomite 3 with-
out dorsal hook or special process.
DESCRIPTION.?Rostrum fully developed, uncon-
stricted. Fuzz on article 1 of antenna 1 in male
absent, calceoli on male primary flagellum of an-
tenna 1 present. Calceoli on male antenna 2 un-
known. Prebuccal parts ordinary, extended forward,
NUMBER 245 41
poorly separated from each other, upper lip domi-
nant. Right lacinia mobilis simple, flabellate; article
1 of mandibular palp short, palp thick, apex of
article 3 oblique, article 2 with outer setae. Lower
lip lacking cones. Outer plate of maxilla 1 with 11
spines (one spine especially thickened). Inner plates
of maxilliped thick, ordinarily setose. Gills present
on coxae 2-7. Coxae 2-4 without special antero-
dorsal humps. All posterior spines on article 6 of
pereopods 1-2 thin and stiff, midapical spine pres-
ent. Article 2 of pereopod 5 with facial setae in
bundles. Peduncle of uropod 1 with dorsolateral
spines widely spread, medial spines widely spread;
peduncle of uropod 2 with medial spines widely
spread. Peduncle of uropod 3 bearing extra sub-
apical setae or spines. Telson with midlateral setules
on each side highly proximal, reduced to one mem-
ber on each side.
TYPE-SPECIES.?Urothoe pinguis Haswell, 1880a
[raonotypy].
COMPOSITION.?Unique.
RELATIONSHIP.?Pontharpinia is considered to be
the most primitive known phoxocephalid even
though it possesses many advanced attributes. Like
Phoxocephalus and Leptophoxoides it bears fully
triturative molars and enlarged gnathopod 2, two
characteristics assumed to be primitive, but unlike
the more advanced Phoxocephalus and Leptophox-
oides it has more setae on the inner plate of maxilla
1, continuously spinose rami on uropods 1-2, and
facial setae on article 2 of pereopod 5, all characters
which might be marks of relationship to gammaroid-
haustorioid ancestors.
Other characters not necessarily confined to Pon-
tharpinia but assumed to be primitive or basic to
the phoxocephalid stock are: dispersed and numer-
ous apical setae on the inner plate of the maxilliped
(none of these formed into short stubby spines); dis-
persed facial setae on article 3 of the maxillipedal
palp; fully developed rostrum; presence of posterior
setae on article 2 of pereopods 3-4 and on coxa 4;
setose dactyl of pereopod 5; broad articles of pereo-
pods 3-4; fully setose maxilla 2; setal brushes on
the telson; widely spread ventral setae on article 2
of antenna 1; short article 2 of antenna 1; widely
spread ventral setae on epimera 1-2; ventral seta-
tion on urosomite 1; subapical setae on peduncle of
uropod 3; outer setosity on palp article 2 of man-
dible; widely spread medial spinosity on peduncles
of uropods 1-2; more than 2 apical setae on article
2 of outer ramus on uropod 3; absence of postero-
proximal setae or spines on article 5 of pereopods
1-2; presence of an inner tooth on the dactyls of
pereopods 1-2; and finally, but very importantly,
the fully flabellate right lacinia mobilis lacking a
proximally fused raker spine.
Plesiomorphic characters in Pontharpinia are
considered to be: short and almost cryptic article 5
on gnathopod 2; weakly ensiform antenna 2; ante-
rior setae on coxa 1; obsolete basofacial setae on
uropod 1; basal setules of telson reduced to one on
each lobe.
Characters of unknown evolutionary position are:
special peduncular spine on uropod 1; absence of
posterior seta on epimera 1-2; absence of double
spine rows on the inner ramus of uropod 1; absence
of lateral spination on the urosome; short and im-
mersed apical nail on the maxillipedal dactyl; un-
equal flagella of antenna 1; lack of anterodorsal
humps on coxae 1-4; absence of facial setae on
coxae 6-7; absence of cones on the lower lip; almost
fully amalgamated upper lip and epistome with up-
per lip dominant; and diverse sets of facial spines
on article 4 of antenna 2.
Pontharpinia pinguis (Haswell)
FIGURES 8-10
Urothoe pinguis Haswell, 1880:325-326, pL 19: fig. 2; 1882:
240-241.
Pontharpinia pinguis.Stebbing, 1906:146 [in part].
Paraphoxus pinguis.?J. L. Barnard, 1960:274-277, pL 44.
Not Pontharpinia pinguis.?Stebbing, 1897:33-34, pi. 9B;
1910:635 [= Tipimegus stebbingi].
DESCRIPTION OF FEMALE.?Head about 24 percent
of total body length, greatest width about 55
percent of length, rostrum unconstricted, narrow,
elongate, strongly exceeding apex of article 3 on an-
tenna 1. Eyes small, either stained or clear of pig-
ment. Article 1 of peduncle on antenna 1 about 1.7
times as long as wide, about twice as wide as article
2, ventral margin with about 11 setules, produced
dorsal apex with one setule, article 2 about 0.4
times as long as article 1, with apicoventral cycle of
14-15 setae, primary fiagellum with 12 articles,
about 0.85 times as long as peduncle, first article
elongate, bearing sparse short aesthetascs, accessory
flagellum with 11-12 articles. Spine formula on
article 4 of antenna 2 = 0-15-10-1 or 0-11-9-5 and
6 special apical spines (Figure 8: fA2), dorsal margin
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 8.?Pontharpinia pinguis (Haswell), female "a," 12.2 mm (c ? female "c," 13.2 mm; /
female "t," 126 mm).
NUMBER 245
FIGURE 9.?Pontharpinia pinguis (Haswell), female "a," 12.2 mm (/ ?= female "f," 12.60 mm).
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
?
FIGURE 10.?Pontharpinia pinguis (Haswell), female "a," 12.2 mm (c
/ = female "t," 12.6 mm).
female "c," 135 mm;
with notches each bearing 4-5 setae and one spine,
ventral margin with numerous long setae and
groups of short to medium spines, no ventrodistal
spine, article 5 about 0.65 times as long as article 4,
facial spine formula = 0, dorsal margin bearing
group of short setae, ventral margin with numerous
setae, 2-3 ventrodistal short to medium spines, fla-
gellum about 0.7 times as long as articles 4?5 of
peduncle combined, with 13 articles. Mandibles
with medium palpar hump; right incisor with 3
teeth; left incisor with 3 humps in one branch;
right lacinia mobilis flabellate, denticulate; left la-
cinia mobilis with 5 teeth; right and left rakers 10;
molars subcylindrical, triturative; palp article 1
short, article 2 with one long inner apical seta and
14 other long and short inner setae, and 13 outer
setae, article 3 about 1.1 times as long as article 2,
oblique apex with 19 spine-setae, basofacial form-
NUMBER 245 45
ula = 0-5. Inner plate of maxilla 1 especially large,
broad, bearing 3 apical and 3 lateral and facial
short setae, palp article 2 with 10 apical-lateral-
medial marginal spines and submarginal setae. In-
ner plate of maxilla 2 narrower and shorter than
outer, outer with 7 apicolateral setae, inner with 5-6
medial setae. Inner plate of maxilliped with 4 large
thick apical spines, 5 apicofacial setae, 4 medial
setae; outer plate with 9 medial and apical spines,
7 apicolateral setae; palp articles 1-2 lacking apico-
lateral seta, medial margin of article 2 strongly
setose, article 3 with about 20 facial setae, pair of
lateral setae, nail of article 4 short, almost im-
mersed, with 2 accessory setules. Coxa 1 expanded
distally, anterior margin weakly convex, main ven-
tral setae of coxae 1-4 = 11-14-14-37, coxa 1 with
extra set of 5 medial setae, with brush of antero-
ventral setae, posteriormost seta of coxae 1-3
slightly shortened or as long as others, anterior and
posterior margins of coxa 4 almost parallel, poste-
rior margin almost straight, posterodorsal corner
rounded, posterodorsal margin short, undulant,
width-length ratio of coxa 4 = 2:3. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1_2 = 14-22-21-15, short anteriors = 5-54(+ 10
facial)-5-4, long anterior = 24(+ 19 facial}-5-25-7;
gnathopods enlarged, especially gnathopod 2, width
ratios of articles 5-6 on gnathopods 1-2 = 23:38
and 26:59, length ratios = 46:62 and 44:80, palmar
humps small and large respectively, palms strongly
oblique, article 5 of gnathopod 1 ovate, posterior
margin rounded, article 5 of gnathopod 2 short,
cryptic, triangular, almost lobate. Pereopods 1-2
similar, lacking combs medially, facial setae formula
on article 4 = 6 and 6, on article 5 = 10-11 and 11,
main spine of article 5 extending to M. 55 on
article 6, article 5 lacking proximoposterior arma-
ment with dense distal brush of setal-spines, spine
formula of article 6 = 12+14 or 12+13 plus large
middistal spine, spines especially thin, acclivity on
inner margin of dactyls of pereopods 1-2 sharp,
produced as tooth, emergent setule short, midfacial
pluseta short, highly distad. Coxae 5-7 posteroven-
tral seta formula = 22-(26+)-20. Only article 4 of
pereopods 3-4 broad, facial spine rows dense, facial
ridge formula on article 2 of pereopods 3-5 = 1-1-2,
width ratios of articles 2, 4, 5, 6 of pereopod 3 =
57:60:30:14, of pereopod 4 = 65:60:32:10, of pereo-
pod 5 = 72:13:12:6, length ratios of pereopod 3 =
92:54:45:47, of pereopod 4 = 76:48:45:64, of pereo-
pod 5 = 103:34:26:23, article 2 of pereopod 5 ex-
ceeding middle of article 5, heavily setose ventrally
and facially, article 2 of pereopod 3 tapering proxi-
mally, posterior margins of article 2 on pereopods
$-4 densely setose, setae short, dactyls elongate,
dactyl of pereopod 5 with stiff inner setae, medial
apex of article 6 finely combed, bearing 3 short
digital processes. Posteroventral corner of epimeron
1 rounded, posterior margin convex, anteroventral
margin with brush of 6 medium setae, ventral face
and margin with 8 medium to long setae; postero-
ventral corner of epimeron 2 rounded, posterior
margin convex, facial setae = 19, above and below
2 ridges; posteroventral corner of epimeron 3 with
large tooth, posterior margin straight, setose, ven-
tral margin naked, face with 2-3 oblique rows of
1-6-13 or 2-3-2-12 (etc.) setae from posterior mar-
gin to middle. Urosomite 1 with 2 ventral setae at
base of uropod 1, 3 brushes of midventral setae
from proximal to distal = 10-11, 4-7, and 10-11,
articulation line complete; urosomite 3 protuberant
dorsally, with 8 basal setae on each side. Rami of
uropods 1-2 with articulate small apical nails and
accessory nails; continuously spinose outer ramus
of uropod 1 with 13-18 dorsal spines (counts here
including accessory nails), in rows of 12 and 6 or 13
and 5, inner ramus with 7; outer ramus of uropod 2
with 10-16 dorsal spines, inner with 6, some spines
on outer rami paired; peduncle of uropod 1 with
6-8 apicolateral spines and setae, no basofacials,
medially with 11-12 marginal setae and spines,
some in groups, plus apical enlarged special spine;
peduncle of uropod 2 with 5-6 dorsal spines, me-
dially with 5 spines widely spread; apicolateral
corner of peduncle on uropods 1-2 lacking comb.
Peduncle of uropod 3 with 5 ventrolateral spines,
dorsally with lateral spine and setule, medial spine
and setule, rami masculine, inner extending to M.
100+ on article 1 of outer ramus, apex with 2 setae,
medial and lateral margins setose, article 2 of outer
ramus elongate, 0.37, bearing 3 short to medium
setae and one spine, medial margin of article 1
setose, lateral margin with 3 acclivities, spine form-
ula = 4-4-3-2, setal formula = 4-4-4?1. Telson un-
usual, length-width ratio =1:1, almost fully cleft,
each apex narrow, almost sharp, lateral acclivity
weak, bearing short lateral setule, spine next medial
greatly longer than setule, each lobe with proximo-
lateral arcuate brush of 15-16 long setae plus one
basodorsal plusetule. Cuticle with ordinary bulbar
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
setules of varying sizes mixed with pipes, surface
bearing fine striations in form of linear fingerprint
pattern, emergent setules branched.
OBSERVATIONS (female).?Ventral crescent and lat-
eral brush of setae on article 2 of antenna 1 sepa-
rated by gap filled with penicillate setules (not
shown on illustration); some long setae on article 2
of gnathopods and pereopods 1-2 paired, especially
posteriorly on gnathopods 1-2 and pereopod 1, and
anteriorly on gnathopod 1; inner ram us of uropod
1 with 3 basomedial spines (all other phoxocepha-
lids observed have only one basofacial spine, seta,
or setule); coxae 5-6 with or without smooth gaps
between setae on posterior margin; coxa 1 bearing
anteromedial facial setal groups, also with or with-
out supernumerary posteroventral setal group on
medial face; apical notch on article 2 of outer
ramus on uropod 3 filled with 1-2 scales.
DESCRIPTION OF MALE.?No advanced males in
collections. Males at hand marked by penial proc-
esses and one large calceolus each on articles 1-3 of
primary flagellum on antenna 1, medial margin of
article 1 on antenna 1 lacking fuzz, no other differ-
entiation on antenna 2 or uropod 3, dactyl of
pereopod 5 flattened, blade-like.
ILLUSTRATIONS.?Pereopod 4 magnified only 84
percent as much as pereopods 3 and 5; black space
on illustration of pereopod 1 at juncture between
articles 3 and 4 representing open space; six special
spines of article 4 on antenna 2 shown only as cir-
cles on Figure 8: fA2.
LECTOTYPE (here examined and designated).?
AM G.5406, 2 slides with 3 cover slips total, bearing
one each of pereopod 4, pereopod 5, uropod 1,
uropod 2, epimera 1-3, and some pleopods, suffi-
cient to recognize as Pontharpinia pinguis; labeled
as "Holotype" but probably not selected by Haswell.
TYPE-LOCALITY.?As published, not on slides,
Bondi, New South Wales, Australia, cast on a beach
during a storm.
VOUCHER MATERIAL.?CPBS 35S/5: female "a,"
12:2 mm (illus.). CPBS 35S/4: female "c," 13.2 mm
(illus.); female "f," 12.60 mm (illus.); male "m,"
10.76 mm.
MATERIAL.?CPBS, 36 samples from 13 stations
(118); PPBES, 5 samples from 2 stations (7); SBS,
3 samples (5); VVAM, one sample (1).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 2-16.5 m, sand, muddy sand, shell. New
South Wales: Bondi Beach (type-locality); off Jib-
bon Point, 40 m; and off Port Stephens, near
Newcastle, 18-22 m, sand.
TIPIMEGINAE
DIAGNOSIS.?Article 2 of antenna 1 ordinary to
elongate. Mandibular molar enlarged but pseudo-
triturative, bearing a special pattern of 3-4 mostly
fused spines. Palp of maxilla 1 biarticulate. Setation
on maxilla 2 ordinary. Gnathopod 2 as small as
gnathopod 1. Article 2 of pereopod 3 of broad
form, not tapering d is tally; pereopod 5 with en-
larged article 3.
SPECIAL CHARACTERS.?Gnathopods of special tri-
chophoxid form, article 5 elongate, article 6 thin,
mitellid, heavily setose anteriorly; uropod 1 with
ventral spike.
DESCRIPTION.?Article 5 of antenna 2 usually re-
duced in size. Epimeron 3 of nonrounded classifica-
tion. Apices of peduncles on uropods 1-2 combed.
TYPE GENUS.?Tipimegus, new genus.
COMPOSITION?Booranus, new genus; Tricho-
phoxus K. H. Barnard; Waitangi Fincham.
Key to the Genera of Tipimeginae
1. Uropods 1-2 with elongate setae on peduncles and rami, epimeron 3 with large postero-
ventral tooth Waitangi
Uropods 1-2 lacking elongate setae, epimeron 3 lacking large posteroventral tooth 2
2. Telson with lateral spines, urosome with lateral spines, inner ramus of uropod 1 with only
one row of marginal spines Trichophoxus
Telson lacking lateral spines, urosome lacking lateral spines, inner ramus of uropod 1
with 2 rows of marginal spines 3
3. Epistome with large anterior tooth, pereopod 5 with gill, epimeron 3 with grossly developed
facial row of spines Booranm, new genus
Epistome lacking anterior tooth, pereopod 5 lacking gill, epimeron 3 lacking facial spine
row Tipimegus, new genus
NUMBER 245 47
Tipimegus, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female; article 2 of antenna 1
especially elongate, ventral setae widely spread.
Article 1 of antenna 2 weakly ensiform; article 3
with 3 or more setae, setules, and spines; facial
spines on article 4 in 2 or more rows, plus special
apical spines; article 5 especially thin and short.
Right mandibular incisor with 4 teeth; molar not
triturative. large, elongate, conical, then subtrun-
cate and bearing 3-1 special large spines, not bear-
ing fuzz; palpar hump small. Palp of maxilla 1
biarticulate, inner plate with 3-5 setae. Setation of
maxilla 2 ordinary. Inner plate of maxillipeds
ordinary, apex of palp article 3 not or weakly pro-
tuberant, dactyl elongate, apical nail absent. Gills
present on coxae 2-6 only. Gnathopods small, simi-
lar, article 5 elongate, free, without eusirid attach-
ment, palms transverse to chelate, hands elongate,
heavily setose anteriorly, trichophoxid in shape.
Article 5 of pereopods 1-2 lacking setae postero-
proximally. Article 2 of pereopod 3 of broad form;
articles 4?5 of pereopods 3-4 broad; article 2 of
pereopod 3 weakly or strongly setose posteriorly;
pereopod 5 especially small, article 2 strongly setose
ventrally, article 3 greatly enlarged, dactyl normal.
Epimera 1-2 bearing numerous long posterior setae,
with or without midfacial setae above ventral facial
ridge; epimeron 3 ordinary, lacking facial spines.
Urosomite 1 naked laterally, bearing one or more
midventral crescents or bundles of setae, urosomite
3 lacking dorsal hook or special process. Peduncle
of uropod 1 weakly to strongly shortened, with large
apicoventral spike, without special enlarged dorso-
lateral-medial spine, only peduncular apices of
uropod 2 combed, inner ramus of uropod 1 with 2
rows of marginal spines, rami often continuously
spinose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, article 2 of outer ramus carry-
ing 3 medium to long apical setae. Telson ordinary,
with 2-4 apical spines, rarely one spine slightly
lateral.
DESCRIPTION.?Rostrum fully developed, con-
stricted. Fuzz on article 1 of male antenna 1 ab-
sent, calceoli on male primary flagellum of antenna
1 present, but developing late. Calceoli on article 5
of male antenna 2 present, facial spines absent in
female, flagellum in male with calceoli. Prebuccal
parts strongly extended forward, massive, epistome
and upper lip distinct, epistome dominant, lacking
process. Right lacinia mobilis bifid, thin; article 1
of mandibular palp short, palp medium to thick;
apex of article 3 oblique, article 2 with outer setae.
Lower lip lacking cones. Outer plate of maxilla 1
with 11 spines, one spine especially thickened. Inner
plates of maxilliped thick, ordinarily setose. Coxae
2-4 with anterodorsal humps. All posterior spines
on article 6 of pereopods 1-2 thick and stiff, mid-
apical spine especially enlarged. Article 2 of pereo-
pod 5 with facial setae. Peduncle of uropod 1 with
dorsolateral spines confined apically, or weakly
spread, medial spines confined apically; peduncle
of uropod 2 with only 1-3 medial spines confined
apically. Peduncle of uropod 3 lacking extra sub-
apical setae or spines. Telson with ordinary pair of
midlateral or dorsal setules on each side.
OBSERVATIONS.?Some or all of basofacial setae on
article 3 of mandibular palp thick, all confined to
basodorsal notch; palp of maxilla 1 with fully facial
setae; article 2 of pereopod 5 with anterodorsal
bulge; uropod 1 lacking basofacial setae in late
instars; male developing facial spines on article 5
of antenna 2, absent in female. Occasional speci-
mens identified as T. kangulun and T. thalerus
have the elements of the comb on the outer ramus
of uropod 2 fine but elongate, rigid, and even,
thereby suggesting that the usual ragged appear-
ance is a result of damage.
TYPE-SPECIES.?Tipimegus thalerus, new species.
COMPOSITION.?T. dinjerrus, new species; T. kan-
gulun, new species; T. kalk.ro, new species; Para-
phoxus stebbingi J. L. Barnard, 1958; T. species
4N; T. species 5N.
RELATIONSHIP.?Tipimegus, its companion Boo-
ranus, plus Trichophoxus and Waitangi comprise a
group of genera characterized by the unusual molar
representing an intergrade between the fully tritura-
tive molar of Pontharpinia and the reduced, spinose
molar typical of most phoxocephalids. The molar in
this group of genera, however, appears to lie on a
blind evolutionary end since no genus with a sim-
pler molar can be related to this Tipimegus group
of genera because of the unusual pereopod 5 in the
group. Pereopod 5 is miniaturized, its article 3 is
enlarged and somewhat similar in shape to the ap-
parently ancestral situation in Pontharpinia. Article
2 of pereopod 5 bulges anteriorly (except in the
genus Booranus) and has facial setae. Gnathopods
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
1-2, with elongate and free wrists bearing mitellid
hands heavily setose anteriorly, could be desig-
nated "trichophoxid" after one of the genera in the
group. The trichophoxid gnathopod is found in
American phoxocephalids also but not in conjunc-
tion with the unusal pereopod 5 so that it has
apparently evolved at least twice in different evo-
lutionary clusters.
The tipimegin group also has more than 2 setules
on article 3 of antenna 2, reminiscent of Ponthar-
pinia. Tipimegin genera are also coordinate in
most of the following characters: a ventral spike
on uropod 1, posteriorly setose epimera 1-2, almost
or fully obsolete nail on dactyl of maxilliped;
anterodorsal humps on coxae 2-4, elongate to
ordinary article 2 of antenna 1 and short article 5
of antenna 2, with loss of facial spines in female
only, elongation of article 5 on pereopods 1-2 with
proximoposterior margin naked, manifold sets of
dorsal setae on article 4 of antenna 2 in female,
posterior setation on coxa 4, extra apical spine or
seta on article 2 of outer ramus on uropod 3,
slightly ensiform antenna 2, scattered facial setae
on palp article 3 of maxilliped, basofacial setae on
article 3 of mandibular palp confined to a dorsal
notch, presence of outer or facial setae on article
2 of mandibular palp, presence of posterior setae on
article 2 of pereopods 3-4, conspicuous middistal
spine on article 6 of pereopods 1-2, and presence of
supernumerary apical spines on article 4 of antenna
2.
Tipimegin characters shared with Pontharpinia
are: presence of a basodorsal notch on article 3 of
the mandibular palp, presence of more than 2
setules on article 3 of antenna 2, absence of facial
spines on article 5 of antenna 2 in the female,
absence of posteroproximal setae on article 5 of
pereopods 1-2, presence of more than 2 apical setae
on article 2 of the outer ramus on uropod 3, pres-
ence of ventral bundles of setae on the urosome,
absence of basofacial setae on uropod 1, a thin
resemblance in setosity on article 2 of pereopods
3-4, and the weakly ensiform antenna 2.
Tipimegus shares with Pontharpinia but not with
other tipimegins the presence of 2 individual rows
of spines on the inner ramus of uropod 1.
Tipimegus and its allies differ from Pontharpinia
in the presence (usually) of 4 teeth on the right
lacinia mobilis, the special molar, the unusual gnat-
hopods, hereafter called "trichophoxid," on which
the hand is thin, mitelloid, the palms transverse, the
hands heavily armed anteriorly with setae and elon-
gate saber spines, the reduction in si/e of pereopod
5 with enlargement of article 3 and reduction of
article 4, the loss of or complete fusion of the apical
nail on the maxillipedal palp, the loss of a special
dorsomedial spine on the peduncle of uropod 1
and the comcomitant development of a special ven-
tral, almost interramal spine, usually attached to a
process and called a "spike," the usually nonflabel-
late (except IVaitangi) and bifid right lacinia mo-
bilis, the elongation of article 2 on antenna 1, the
elongation of one or more setae on the inner plate
of maxilla 1, the greater breadth, wider truncation,
and more apically confined armament on the palp
of maxilla 1, and the reduction in number of thick
apical spines on the apex of the inner plates of the
maxilliped, generally from 4 down to 2 or fewer.
The midapical spine on article 6 of pereopods 1-2
usually becomes highly enlarged in the tipimegin
group, and the rostrum is strongly narrowed. All
tipimegins appear to share massive prebuccal parts.
Tipimegus differs from Trichophoxus K. H.
Barnard in the absence of lateral spines on the
telson, in the absence of spines on the inner ramus
of uropod 2, in the presence of 2 rows of spines on
the inner ramus of uropod 1, in the apical confine-
ment of medial spines on the peduncle of uropod 2,
in the somewhat narrow spread of the dorsolateral
spines on the peduncle of uropod 1 and the slight
to great shortness of that peduncle. Tipimegus also
has a much larger spike on uropod 1 than does
Trichophoxus, but the latter genus has a much
more greatly enlarged midapical spine on article 6
of pereopods 1-2 than does Tipimegus. Tricho-
phoxus also bears facial setae on article 2 of pereo-
pod 4.
Tipimegus and Booranus are composed of two
groups of closely related tipimegins. Tipimegus
differs from Booranus in the following characters:
absence of a strong oblique row of spines on epi-
meron 3, presence of midfacial setae on epimeron 1,
strong posterior setation on article 2 of pereopod 3,
presence of only 3 setae on the inner plate of
maxilla 1 (as against 4-5), presence of only one
apicomedial spine on the peduncle of uropod 2
(as against 2-3), apical confinement of the dorso-
lateral spines on peduncle of uropod 1, narrow
spread of medial spines on that same peduncle, and
the shortness of that peduncle.
NUMBER 245 49
Booranus forms a weak bridge between Tigimegus
and Trichophoxus in the characters of uropod 1.
The form of spination in tipimegins on article
4 of antenna 2 differs from other phoxocephalids in
that an extra fully apical, often widespread row of
spines occurs; this is quoted first and usually com-
prises 2 to 5 spines. The second row, highly oblique,
is often composed of 2 subgroups, especially notice-
able in Booranus. When the citation 4-2-8-7-6
(example) is cited, the second row has been noted
as being divided into a set of 2 and 8. In subsidiary
descriptions this division of row 2 is often ignored
and the first marginal row is not quoted at all.
In the descriptions the apical nails and accessory
apical nails are included in the dorsal spine counts
on rami of uropods 1-2 in Booranus but not in the
other tipimegin genera. We are uncertain as to the
homology of apical nails in the tipimegin genera;
they are so erect and free that their arrangement is
quite distinct from the situation in the birubiin
group but somewhat similar to conditions in the
harpiniin subfamily.
Key to the Species of Tipimegus (Adults)
(Species "iX cited twice, 7'. stebbingi not included)
1. Comb on outer ramus of uropod 2 conspicuous at X 40 magnification (Figure 23: OR2c),
grossly uneven T. kalkro, new species
Comb on outer ramus of uropod 2 inconspicuous at x 40 magnification, fine and dense,
or comb abraded or absent 2
2. Hands of gnathopods narrow apicaliy (Figure 13: Gl) 3
Hands of gnathopods broad apicaliy (Figure 15: Gl) 4
3. Palm of gnathopod 1 s-shaped, dactyl and article 6 of pereopod 3 elongate, article 6 very
thin Tipimegus sp. 5N
Palm of gnathopod 1 straight, dactyl and article 6 of pereopod 3 not elongate, article 6
stout T. thalerus, new species
4. Setae on coxa 1 spread over more than half of ventral margin, outer ramus of uropod 1
with dorsal spination not continuous evenly to apex T. kangulun, new species
Setae on coxa 1 spread over less than one-fourth of ventral margin, outer ramus of uropod
1 with dorsal spination almost fully continuous to apex 5
5. Palm of gnathopod 1 s-shaped Tipimegus sp. 5N
Palm of gnathopod 1 even 6
6. Epimeron 2 with setae fully in middle of plate, dactyl of pereopod 3 much longer than half
of article 6, outer ramus of uropod 1 with large accessory spines contiguous with nail
Tipimegus sp. 4N
Epimeron 2 lacking setae in middle of plate, dactyl of pereopod 3 much less than half as
long as article 6, outer ramus of uropod 1 with large accessory spines contiguous with
nail T. dmjerrus, new species
Tipimegus thalerus, new species
FIGURES 11-14
DESCRIPTION OF FEMALE.?Head about 23 percent
of total body length, greatest width about 70 per-
cent of length, rostrum constricted, elongate, reach-
ing middle of article 2 on antenna 1. Eyes small,
largely occluded with pigment, ommatidia espe-
cially small. Article 1 on peduncle of antenna 1
about 1.3 times as long as wide, about 2.3 times as
wide as article 2, apicoventral margin with about 10
setules, weakly produced dorsal apex with 2 setules;
article 2 about 1.1 times as long as article 1, with
ventral cycle of 11-12 setae; primary flagellum with
10-13 articles, basal elongate, about 0.65 times as
long as peduncle, bearing several aesthetascs; acces-
sory flagellum almost as long as primary flagellum,
with 10-11 articles. Spine formula on article 4 of
antenna 2 = 2-2-7-^-6 or 2-11-12-8, with one
supernumerary spine, dorsal margin with 4 notches
bearing 4-6 setae each, ventral margin with 6
groups of 1-6 long to medium setae, dense ventro-
distal brush of setae; article 5 about 0.45 times as
long as article 4, facial spine formula = 0, dorsal
margin bearing weak clumps of setae, ventral mar-
gin with 9 unpaired setae and clump of 4-5 long to
medium spine-setae, this row set subfacially; flagel-
lum about 0.85 times as long as articles 4-5 of
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 11.?Tipimegus thalerus, new species, holotype, female "a," 7.7 mm (b ? male "b," 6.20 mm).
NUMBER 245 51
s
FIGURE 12.?Tipimegus thalerus, new species, holotype, female "a," 7.7 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
.
FIGURE IS.?Tipimegus thalerus, new species, holotype, female "a," 7.7 mm (/ ?? juvenile "j,"
6.58 mm; q ? female "q," 8.46 mm).
peduncle combined, with 12 articles. Epistome large
but unproduced anteriorly; upper lip with weak
ventral sinus. Mandibles with weak palpar hump;
right incisor with 4 teeth; left incisor with 4 humps
in 2 branches; right lacinia mobilis bifid, distal
branch much shorter than proximal, subbifid, prox-
imal branch simple, pointed; left lacinia mobilis
with 4 teeth; right rakers 11, left rakers 12; molars
elongate, conical or columnar, each with 4 long to
short spines, none disjunct (as in T. kangulun);
palp article 1 short, article 2 with 2 long inner
apical setae and 5 other shorter inner and facial
setae, article 3 about as long as article 2, oblique
apex with 11 spine-setae, basodorsal formula = 3.
Inner plate of maxilla 1 short, broad, bearing 3
long apical setae, one scarcely more elongate than
others, plumose or not; palp article 2 with 6-8
apicalmedial marginal spines and 7 submarginal
and facial setae. Inner plate of maxilla 2 shorter
and narrower than outer, outer lacking apicolateral
setae, inner with 10 medial setae. Inner plate of
maxilliped with 2 large, thick apical spines, 5
apicofacial setae, 5-6 medial setae; outer plate with
15 medial and apical spines, one apicolateral seta;
palp article 1 lacking apicolateral seta, article 2
with 5 groups of 1-2 apicolateral setae, medial mar-
gin of article 2 moderately setose, article 3 with
nearly 20 facial setae, 7 lateral setae, article 4 with
2 accessory setules, nail absent. Coxa 1 strongly
expanded apically, anterior margin almost straight,
NUMBER 245 53
w
FIGURE 14.?Tipimegus thalerus, new species, holotype, female "a," 7.7 mm (b ? male "b," 650
mm; q ? female "q," 8.46 mm).
main ventral setae of coxae 1-4 = {7-8)-(8-l l)-(10-
ll)-(25-26), posteriormost seta of coxae 1-3 short-
est, anterior and posterior margins of coxa 4 strongly
divergent, posterior margin oblique, almost straight,
posterodorsal corner sharp, posterdorsal margin
short, ordinary, undulant, width-length ratio of
coxa 4 = 1:1. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 7-6-9-6 and 4
facial on gnathopod 1, long anteriors = 14-14-22-3,
short anteriors ? 4-2-3-4, no others. Gnathopods
with thin hands narrowed apically; width ratios of
articles 5-6 on gnathopods 1-2 = 27:23 and 25:21,
length ratios = 63:50 and 62:44; palmar humps very
small, palms transverse; article 5 of gnathopods 1-2
elongate, ovate, posterior margin rounded. Pereo-
pods 1-2 similar; facial setae formula on article
4 = 5 and 7, on article 5 = 4-5 and 5-7; main spine
of article 5 very thin, extending to M. 55 on article
6, article 5 with one extra long distoposterior spine
and numerous setae, proximal margin naked, spine
54 SMITHSONIAN CONTRIBL"! IONS TO ZOOLOGY
formula of article 6 = 7 + 9 and 8 + 9 or 7 + 10
and 8 + 1 0 plus huge middistal spine, several spines
elongate; acclivity on inner margin of dactyls of
pereopods 1-2 sharp, produced as tooth, emergent
setule short, marginal pluseta short, coxae 5-7
posteroventral seta formula = 28-29-25, composed
of marginal and submarginal groups. Articles 4-5 of
pereopods 3?4 broad, facial spine rows dense, facial
ridge formula on article 2 of pereopods 3-5 = 0-1-1,
width ratios of articles 2, 4, 5 / 6 of pereopod 3 =
52:48:43:17, of pereopod 4 = 59:59:42:21, of pereo-
pod 5 = 56:9:8:3; length ratios of pereopod 3 =
71:45:38:32, of pereopod 4 = 68:60:4945, of pereo-
pod 5 = 73:18:15:17; article 2 of pereopod 5 almost
reaching apex of article 5, ventrally setose; medial
apex of article 6 finely combed only on pereopod 5;
dactyls of pereopods 3-4 short. Posteroventral corner
of epimeron 1 rounded-quadrate, posterior margin
weakly convex, setose, anteroventral face with 12 +
short to medium setae, midface with 11 medium
setae; posteroventral corner of epimeron 2 rounded,
posterior margin weakly convex, setose, facial
setae = 1 1 , posterior pairs and triads set vertically;
posteroventral corner of epimeron 3.weakly protub-
erant, posterior margin straight, weakly serrate,
setose, ventral margin with 16 setae mainly in
posterior half, face naked. Urosomite 1 with 3
paired sets of setae, each side with sets of 10-11,
ventrally with sets of 3-4 and 3, articulation line
complete; urosomite 3 protuberant dorsally. Rami
of uropods 1-2 with articulate enlarged sharp apical
nails, outer and inner rami of uropods 1-2 with
accessory nails, continuously spinose; outer ramus
of uropod 1 with 9-10 dorsal spines, inner with 4-5
dorsal and 4 medial; outer ramus of uropod 2 with
6 dorsal spines, inner with 2 dorsal spines; peduncle
of uropod 1 with 2-3 apicolateral spines, no baso-
facial setae, medially with 3 marginal setae and
spines in apical clump; peduncle of uropod 2 with
4 dorsal spines, medially with one small apical spine,
apicolateral corner on peduncle of uropod 2 with
weak comb, rami naked. Peduncle of uropod 3 with
12-13 ventral spines, dorsally with one lateral spine,
one medial spine and setule; rami masculine, inner
extending to M. 100+ on article 1 of outer ramus,
apex with 2 setae, medial and lateral margins setose;
article 2 of outer ramus ordinary, 0.21 as long as
article 1, bearing 2 medium setae and one spine;
.apicomedial margin of article 1 setose, lateral mar-
gin with 5 acclivities, spine formula = 0-2-2-2-2-2
or 2-2-2-2-2, setal formula = I-fMMMM), or none.
Telson ordinary, length-width ratio = 50:53, not
fully cleft, each apex wide, rounded, middle accliv-
ity shallow, lateral acclivity tiny, bearing 1-2 ordi-
nary lateral spines, spine next medial longer, with
1-2 lateral and one medial spines separated by
setule, midlateral setules diverse. Cuticle with en-
larged but sparse bulbur setules, surface bearing fine
denticles, emergent setules especially long, branched.
DESCRIPTION OV MAI K (suhadult, male "b," 6.20
mm).?Eyes enlarged, clear of occluding pigment.
Article I of antenna 1 with 13 ventrodistal setules,
article 2 with 12 setae in ventral spray, primary
Hagellum with 10 articles, long acsthetascs on arti-
cles 4-9, accessory Hagellum with 9 articles. Spine
formula on article 4 of antenna 2 = 2-9-6-fi, on
article 5 1. Hagellum only weakly elongate, with
14 articles, not proliferate, calceoli absent. Setal
formula on coxae 1-4 = 7-8-8-14. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 3-3-3-4, long ameriors = 12 (plus 2 facial)-
9-9-1, short anteriors = 1-4-3-4, no others; hands
of gnathopods even thinner than in adult female.
Setal and spine formulas on pereopods 1-2, article
4 = 5 and 5, article 5 = 5 and 5, article 6 = 6 + 7
plus huge middistal spine, articles 4-5 thinner than
in female. Coxa of pereopod 5 with gills besides
penial processes(l). Setal formula on epimera: epi-
meron 1 facial = 8, anteroventral = 6, epimeron 2
anteroventral = row of 6 only, epimeron 3 ven-
tral = 10. Spine formulas on uropods: peduncle of
uropod 1 = 2 dorsolateral, 3 medial, of uropod 2 =
4 dorsal, one medial; outer ramus of uropod 1 = 9
dorsal, of uropod 2 = 6, inner ramus of uropod
1 = 5 dorsomedial, 4 lateral, of uropod 2 = 2.
Uropod 3 peduncle ventral = 12, article 1 of outer
ramus lateral spines = 1-1-1-2-2-2, setae = 0 -0 -1 -
0-0-0, apicolateral spine on article 2 as in female
but slightly more elongate, as in juveniles. Telson
weakly elongate, each lobe with 3 spines, no den-
ticles. Cuticular setules enlarged as in female.
OBSERVATIONS.?Head of holotype as illustrated
showing only weak pointed lobe, this lobe usually
larger and more pointed in other specimens. Article
3 of antenna 2 with 6 facial spines and/or setae plus
one disjunct setule.
VARIATIONS.?Female "t," 13.7 mm: Possibly
gerontic but brood plates fully developed; proximal
NUMBER 245 55
spine formula on article 4 of antenna 2 = 14-11-9,
flagellum with 13 articles; spike on uropod 1 short,
scarcely exceeding M. 50 on outer ramus, peduncle
of uropod 1 with 3 dorsolateral spines, outer ramus
with 15 spines plus accessory spinule at base of nail,
inner ramus with 8 medial and 5 lateral spines,
outer ramus of uropod 2 with 9 dorsal spines.
Juvenile " j , " 6.58 mm: Pereopods 1-2 drawn
from this specimen, these pereopods of shape iden-
tical to adult female, setal, and spine counts on
pereopods 1-2 article 4 = 5 and 5, article 5 = 4 and
4, article 6 = 6 + 8 and 7 + 8 plus huge middistal
spine; uropod 1 with 3 dorsolateral peduncular
spines; article 2 on outer ramus of uropod 3 with
apicolateral spine as in adult.
ILLUSTRATIONS.?Most of pereopod 5 hidden from
lateral view by pereopod 4 in body illustration; an-
tenna 2 projecting fully lateral but flattened in
body illustration; in enlarged views of uropods 1-2
latter enlarged more than former; female pereopod
5 shown in 3 views, small view at same magnifica-
tion as pereopods 3-4, enlarged views from lateral
and medial, note aberrancy of lateral view article 2.
HOLOTYPE.?NMV, female "a," 7.7 mm.
TYPE-LOCALITY.?CPBS 35S, 3 Aug 1972, Western
Port, Victoria, Australia, 14 m, sand.
VOUCHER MATERIAL.?CPBS 25N/1: male "b,"
6.20 mm (illus.); female "r," 13.0 mm. CPBS 26N/4 :
female "q," 8.46 mm (illus.); juvenile " j , " 6.58 mm
(illus.). CPBS 32E/1171: gerontic female "t," 13.7
mm.
RELATIONSHIP.?As the type-species of Tipimegus
this forms the model to which following species are
compared. The main taxonomic characters of T.
thalerus are as follows: continuous spination on the
outer ramus of uropod 1, short dactyls of pereo-
pods 3-4, highly enlarged cuticular setules, absence
of combs on rami of uropods 1-2, apically narrow
hands of gnathopods, narrow spread of ventral setae
on coxae 1-3, and the transformation of the lateral
(number 3) seta on article 2 of the outer ramus on
uropod 3 into a stiff spine.
MATERIAL.?CPBS, 10 samples from 7 stations
(14); PPBES, 5 samples from one station (6); AM,
one sample (1); SBS, one sample (1).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 9-14 m, sand. New South Wales: off
Manly Beach, Sydney, 6 m, sand; off Malabar, 31 m,
sand and shell.
Tipimegus dinjerrus, new species
FIGURE 15
DESCRIPTION OF FEMALE.?Head about 22 percent
of total body length, greatest width about 75 per-
cent of length, rostrum constricted, narrow, exceed-
ing middle of article 2 on antenna 1. Eyes medium,
clear of pigment, ommatidia sparse and scattered.
Article 1 on peduncle of antenna 1 about 1.2 times
as long as wide, about 2.3 times as wide as article 2,
ventral margin with about 8 setules, weakly unpro-
duced dorsal apex with 3 setules, article 2 about
1.05 times as long as article 1, with apicoventral
crescent of 11 setae; primary flagellum with 11
articles, about 0.6 times as long as peduncle, bear-
ing several long aesthetascs (articles 3-10), accessory
flagellum about as long as primary flagellum, with
10 articles. Spine formula on article 4 of antenna
2 = 2-2-9-12-7, with 2 supernumerary spines,
dorsal margin setose, ventral margin with 8 groups
of 2-4 long to medium setae, dense ventrodistal
brush of setae; article 5 about 0.55 times as long as
article 4, facial spine formula = 0, dorsal margin
naked, ventral margin with 5 sets of one seta each, 5
ventrodistal long to medium spine-setae, this row
set facially; flagellum about 0.9 times as long as
articles 4-5 of peduncle combined, with 12 articles.
Mandibles with weak to medium palpar hump;
right incisor with 4 teeth, left incisor with 5 humps
in 2 branches; right lacinia mobilis bifid, distal
branch shorter than proximal, subbifid, proximal
branch simple, pointed, lacking marginal denticles;
left lacinia mobilis with 4 teeth; right rakers 10 plus
one rudimentary, left rakers 11; molars elongate,
conical or columnar, each with one long apical
spine, one short apical spine, one medium spine on
side with strong nonarticulate partner; palp article
1 short, article 2 with 2 long to medium inner apical
setae and 3 other shorter inner setae, and 2 outer
setae, article 3 about 0.9 times as long as article 2,
oblique apex with 12 spine-setae, basodorsal for-
mula = 2. Inner plate of maxilla 1 ordinary,
bearing one long subapical pluseta, 2 apicolateral
shorter setae; palp article 2 with 6 apical spines and
7 submarginal and facial setae. Inner plate of max-
illa 2 shorter and much narrower than outer,
outer lacking apicolateral setae, inner with 9 medial
setae. Inner plate of maxilliped with 2 large, thick
apical spines, 7 apicofacial setae, 4 medial setae;
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 15.?Tipimegus dinjerrus, new species, holotype, female "w," 8.1S mm (v ? male "v,"
7.85 mm).
outer plate with 14 medial and apical spines, one
apicolateral seta; palp article 1 lacking apicolateral
seta, article 2 with 4 groups of one each apicolateral
setae, medial margin of article 2 moderately setose,
article 3 with 14-16 facial setae, 6 lateral setae, nail
of article 4 absent, with 3 accessory setules. Coxa 1
strongly expanded distally, anterior margin almost
straight; main ventral setae of coxae 1-4 = 9-11-
10-25, posterionnost seta of coxae 1-3 shortest;
anterior and posterior margins of coxa 4 strongly
divergent, posterior margin oblique, almost straight,
posterodorsal corner sharp, posterodorsal margin
ordinary, almost straight, width-length ratio of coxa
4 = 11:13. Long posterior setae of article 2 on
gnathopods 1-2 and pereopods 1-2 = 6-6-7-6.
long anteriors = 15(+ 6 facial)-! 1-20-2, short an-
teriors = 2-4-7-5, no others; gnathopods with
thin hands broad apically, width ratios of arti-
cles 5-6 on gnathopods 1-2 = 29:26 and 27:25,
length ratios = 65:52 and 65:45, palmar humps
very small, palms transverse, article 5 of gnathopods
1-2 elongate, ovate, posterior margins rounded-flat
NUMBER 245 57
Pereopod 2 stouter than pereopod 1, especially arti-
cle 4; facial setae formula on article 4 = 6 and 6.
on article 5 = 5 and 6, main spine of article 5 ex-
tending to M. 75 on article 6, spine formula of arti-
cle 6 = 8 4- 11 and 8 + 1 1 plus huge middistal spine;
acclivity on inner margin of dactyls of pereopods
1-2 sharp, produced as tooth, emergent setule short,
midposterior pluseta short. Coxae 5-7 posteroven-
tral seta formula = 28-26-30, composed of margi-
nal and facial groups. Articles 4-5 of pereopods 3-4
broad, facial spine rows dense; facial ridge formula
on article 2 of pereopods 3-5 = 0-1-1; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 52:48:45:17,
of pereopod 4 = 62:57:37=19, of pereopod 5 =
64:10:9:5; length ratios of pereopod 3 = 74:42:
44:30, of pereopod 4 = 70:60:46:48, of pereopod
5 = 75:20:16:20; article 2 of pereopod 5 exceed-
ing middle of article 5, ventrally setose, with 7
facial setae (long) and several short setae; medial
apex of article 6 only on pereopod 5 finely combed;
dactyls of pereopods 3-4 short. Posteroventral
corner of epimeron 1 rounded, posterior margin
wealJy convex, setose, anteroventral face with 8
short to medium setae, midface with 5 medium
setae; posteroventral corner of epimeron 2 rounded,
posterior margin weakly convex, setose, facial setae
= 10, in rows of 7 and 3; posteroventral corner of
epimeron 3 weakly protuberant, posterior margin
straight, weakly serrate, setose, ventral margin with
11 setae mainly in posterior half, 4 of these setae
in 2 pairs, face with occasional submarginal seta.
Urosomite 1 with 4 paired sets of setae, each side
with sets of 5, ventrally with sets of 4, 3, and (2^),
articulation line complete, urosomite 1 protuberant
dorsally. Rami of uropods 1-2 with articulate
enlarged sharp apical nails; outer and inner rami
of uropod 2 with accessory nails, continuously
spinose; outer ramus of uropod 1 with 9 dorsal
spines, almost continuously spinose, with small
asymmetricity, inner with 6 dorsal and 3 lateral
spines; outer ramus of uropod 2 with 7 dorsal
spines, inner with one spine (= accessory); peduncle
of uropod 1 with 3-4 apicolateral spines, medially
with 3 marginal spines, peduncle of uropod 2 with
5 dorsal spines, medially with one small apical
spine, apicolateral corners of peduncles on uropods
1-2 and rami lacking comb. Peduncle of uropod 3
with 17 ventral spines, dorsally lacking lateral spine
(merged with ventral spines), one medial spine and
setule; rami subraasculine, inner extending to M.
100 on article 1 of outer ramus, apex with 3 setae,
medial and lateral margins setose; article 2 of outer
ramus short, 0.14, bearing 3 medium to long setae,
lateralmost being medium unplumose spine, apico-
medial margin of article 1 setose, lateral margin
with 3 acclivities, spine formula = 2-2-2-2, setal
formula = 0-0-0-1 or 0. Telson ordinary, length-
width ratio = 25:28, not fully cleft, each apex of
medium width, rounded, undulant, lateral acclivity
shallow, weak, with lateral and medial spines
separated by setule; midlateral setules diverse.
Cuticle with ordinary bulbar setules mixed with
pipes, surface bearing fine denticles, emergent
setules branched.
OBSERVATION (female).?Article 3 of antenna 2
with 6 facial spines and setae plus one disjunct set-
ule; rostrum much narrower than in T. thalerus.
DESCRIPTION OF MALE (subadult).?Eyes scarely
enlarged; primary flagellum of antenna 1 with 10
articles, bearing long aesthetascs on articles 2-9;
dorsal margin of article 4 on antenna 2 with male
fuzz, spine formula = 2-10-9-5, of article 5 = 2,
article 5 with 5 sets of dorsal male setae, flagellum
weakly elongate, with 15 articles, calceoli absent;
basofacial setal formula on article 3 of mandibular
palp = 3, inner margin with 2 medium sized acces-
sory setae; article 4 of pereopods 1-2 much thinner
than in female; pereopods 3-5 and epimera similar
to those of female; see illustrations for spines on
uropods 1-2; spine formula on article 1 of outer
ramus on uropod 3 = 1-2-2-2-2, setae = 1?1?1?1?
1, setae otherwise scarcely more dense than in
female, peduncle with 12 ventral spines, one dor-
solateral spine; telson as in female but with sparse
mediobasal denticles on each lobe.
ILLUSTRATIONS.?All parts, except those few illus-
trated and except for proportions and spine counts
mentioned in description, like those of Tipimegus
thalerus, especially epistome, mouth parts, pereo-
pods 1-5, dactyls of pereopods 3-4, uropod 3, telson,
especially article 5 of pereopod 5 and setal dis-
tribution on coxae 2-3.
HOLOTYPE.?WAM, female "w," 8.13 mm.
TYPE-LOCALITY.?WAM 416-73, 24 Jul 1943,
off Jibbon Point, New South Wales, Australia, 40 m.
VOUCHER MATERIAL.?The type-locality: male
"v," 7.85 mm (illus.) and 2 similar males.
RELATIONSHIP.?This species appears closest to T.
thalerus mainly because so many minor proportions,
such as those of pereopods 3-5, and the general large
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
size of and setation densities on epimera and coxae
1-3, resemble that species rather than T. kangulun.
Nevertheless T. dinjerrus differs from T. thalerus in
characters it shares with T. kangulun, such as the
small cuticular setules, almost continuous spina-
tion on the outer ramus of uropod 1, and the broad
apices on the hand of the gnathopods. From T.
kangulun, T. dinjerrus differs mainly in the short
dactyls of pereopods 3-4 and the distinctiveness of
the lateral spine on article 2 of the outer ramus
on uropod 3. Only one female and 4 subadult males
of T. dinjerrus are known; thus a wider variety of
material is required to determine whether or not
this species is valid.
MATERIAL.?WAM, one sample (5).
DISTRIBUTION.?New South Wales, off Jibbon
Point, 40 m.
Tipimegus kangulun, new species
FIGURES 16-19
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total length, greatest width about 77 percent
of length, rostrum constricted, narrow, exceeding
middle of article 2 on antenna 1. Eyes small, clear
of pigment or darkly stained. Article 1 of peduncle
of antenna 1 about 0.8 times as long as wide, about
2.8 times as wide as article 2, ventral margin with
about 7 setules, weakly produced dorsal apex with
3 setules, article 2 about 0.9 times as long as article
1, with ventral crescent of 13-14 setae; primary
flagellum with 9 articles, about 0.55 times as long
as peduncle, bearing several long aesthetascs; acces-
sory flagellum almost as long as primary flagellum,
with 8-9 articles. Spine formula on article 4 of an-
tenna 2 = (2-3)-2-l 1-10-7 + 2-3 supernumerary
spines, dorsal margin setose, ventral margin with 8-9
groups of 2-5 long to short setae, dense ventrodistal
brush of setae; article 5 about 0.5 times as long as
article 4, facial spine formula = 0, dorsal margin
naked, ventral margin with 6 unpaired setae, 6 ven-
trodistal long to medium spine-setae, this row set fa-
cially; flagellum about 0.75 times as long as articles
4-5 of peduncle combined, with 10 articles. Man-
dibles with weak to medium palpar hump; right
incisor with 4 teeth; left incisor with 5 humps in 2
branches; right lacinia mobilis bifid, distal branch
shorter than proximal, subbifid proximal branch
simple, pointed, with marginal denticles; left
lacinia mobilis with 1 teeth; right rakers 10 plus
one rudimentary; left rakers II; molars elongate,
conical or columnar, each with one long apical
spine, one short apical spine, one medium spine on
side with vestigial nonarticulate partner; palp arti-
cle 1 short, article 2 with 4-5 medium inner apical
setae, 3 other shorter inner setae and 2-3 outer setae,
article 3 about 0.9 times as long as article 2, oblique
apex with 14 spine-setae, basodorsal formula = 3.
Inner plate of maxilla 1 ordinary, bearing one long
subapical pluseta, 2 apicolateral shorter setae; palp
article 2 with 6-7 apical spines and 9 submarginal
and facial setae. Inner plate of maxilla 2 shorter
and much narrower than outer, outer lacking apico-
lateral setae, inner with 9-10 medial setae. Inner
plate of maxilliped with 2 large, thick apical spines,
6 apicofacial setae. 7 medial setae, outer plate with
14 medial and apical spines, no apicolateral setae;
palp article 1 lacking apicolateral seta, article 2 with
3 groups of 1-2 apicolateral setae, medial margin of
article 2 moderately setose, article 3 with 11 facial
setae, 4 lateral setae, article 4 with 3 apical accessory
setules, nail absent. Coxa 1 strongly expanded dis-
tally, anterior margin almost straight; main ventral
setae of coxae 1-4-14-16-15-32, posterior-most seta
of coxae 1-3 shortest; anterior and posterior mar-
gins of coxa 4 strongly divergent, posterior margin
oblique, almost straight, posterodorsal corner sharp,
posterodorsal margin ordinary, almost straight,
width-length ratio of coxa 4 = 27:29. Long poste-
rior setae on article 2 of gnathopods 1-2 and pereo-
pods 1-2 = 9-7-8-5, long anteriors = 20(+ 6 fac-
ial)-! 9-(15-20)-5, short anteriors = 2-3-2-5, no
others. Gnathopods with thin hands broad apically,
width ratios of articles 5-6 on gnathopods 1-2
= 30:26 and 27:25, length ratios = 63:50 and
63:45; palmar humps very small, palms transverse;
article 5 of gnathopods 1-2 elongate, ovate, posterior
margins rounded-flat. Pereopod 2 stouter than pe-
reopod 1, especially article 4; facial setae formula on
article 4 = 10-11 and 10, on article 5 = 5 and 5,
main spine of article 5 extending to M. 75 on article
6, spine formula of article 6 = 6 + 9 or 7 + 9 plus
huge middistal spine; acclivity on inner margin of
dactyls of pereopods 1-2 sharp, produced as tooth,
emergent setule short, midposterior pluseta short.
Coxae 5-7 posteroventral seta formula = 32-32-20,
composed of marginal and facial groups. Articles
4-5 of pereopods 3-4 broad, facial spine rows dense;
facial ridge formula on article 2 of pereopods 3-5 =
NUMBER 245
s
FIGURE 16.?Tipimegus kangulun, new species, holotype, female "n," 7.60 mm (u - male
"u," 6.72 mm; tv ? female "w," 7.20 mm; y ? male "y," 7.10 mm).
60
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 17.?Tipimegus kangulun, new species, holotype, female "n," 7.60 mm (y ?? male "y,1
7.10 mm).
NUMBER 245 61
FIGURE 18.?Tipimegus kangulun, new species, holotype, female "n," 7.60 mm (j ? juvenile "j,1
4.37 mm; u ? male "u," 6.72 mm; y ? male "y," 7.10 mm; z ? female "z," 11.25 mm).
0-1-1; width ratios of articles 2, 4, 5, 6 of pereopod
3 = 53:48:42:18, of pereopod 4 = 63:59:44:20, of
pereopod 5 = 55:11:10:5; length ratios of pereopod
3 = 72:44:44:34, of pereopod 4 = 75:62:47:45, of
pereopod 5 = 80:23:19:22; article 2 of pereopod 5
reaching middle of article 5, ventrally setose;
medial apex on article 6 of pereopods 3 and 5 finely
combed; dactyls of pereopods 3-4 elongate. Postero-
ventral corner of epimeron 1 rounded, posterior
margin convex, setose, anteroventral face with 15
short to medium setae, midface with 11 medium
set^e; posteroventral corner of epimeron 2 rounded,
weakly protuberant, posterior margin convex,
setose, facial setae = 1 5 + 2 midfacial, pairs and
triads set vertically; posteroventral corner of epi-
meron 3 weakly protuberant, posterior margin
straight, weakly serrate, setose, ventral margin with
16 setae mainly in posterior half, face with occa-
sional sub marginal seta. Urosomite 1 with 4 paired
sets of setae, each side with sets of 8 and 2, ven-
trally with sets of 3 and 2, articulation line com-
plete; urosomite 3 protuberant dorsally. Rami of
uropods 1-2 with articulate enlarged sharp apical
nails, rami of uropod 2 with accessory nails, con-
tinuously spinose; outer ramus of uropod 1 with 9
dorsal spines, irregularly gapped apically, inner
with 4 dorsal and 3 lateral (actually medial and
lateral) spines; outer ramus of uropod 2 with 6-7
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
uW
FIGURE 19.?Tipimegus kangulun, new species, holotype, female "n," 7.60 mm (/ ? juvenile "j ,"
4.37 mm; u ? male "u," 6.72 mm; w - female "w," 7.20 mm; y ? male "y," 7.10 mm; z ?
female "z," 1125 mm).
NUMBER 245 63
dorsal spines, inner with 2-3 dorsal spines;
peduncle of uropod 1 with 3 apicolateral spines,
medially with 4 spines; peduncle of uropod 2 with
4-5 dorsal spines, medially with one small apical
spine; outer rami of uropods 1-2 and apicolateral
corner of peduncle on uropod 2 with weak even
comb. Peduncle of uropod 3 with 13-14 ventral
spines, dorsally with one lateral spine, one medial
spine and setule; rami submasculine, inner extend-
ing to M. 100+ on article 1 of outer ramus, apex
with 2 setae, medial and lateral margins setose,
article 2 of outer ramus short, 0.16, bearing 3
medium to long setae, medial margin of article 1
setose, lateral margin with 3-4 acclivities, spine
formula = 2-2-2-2-( + 2), or 1-2-2-2-2, setal form-
ula = 0. Telson ordinary, length-width ratio = 25:
29 or 26:27, not fully cleft, each apex of medium
width, rounded, undulant, lateral acclivity shallow,
weak, with lateral and medial spines separated by
setule, occasionally lateral spine position paired,
midlateral setules diverse. Cuticle with ordinary bul-
bar setules, surface punctate or bearing fine stria-
tions in form of linear fingerprint pattern, emergent
setules branched.
OBSERVATIONS (female).?Unspined subapex of
outer ramus of uropod 1 often with spine vestige
subapically, juveniles as small as 3.7 mm also lack-
ing spines apically or juvenile "j," 4.37 mm, with sub-
apical spine and second spinule on face of nail, this
spinule also present in superfemale; inner ramus of
uropod 1 in juvenile with pair of accessory apical
spines; protuberant base of spike on peduncle of
uropod 1 becoming elongate in terminal adults.
VARIATIONS (female).?Female "w," 7.20 mm:
Article 5 of pereopod 3 with 2 rows of posterior
facial spines, article 6 of pereopod 4 with 4 rows of
posterior facial spines; spine formula on outer
ramus of uropod 3 = 2-3-2-2-2.
Superfemale "z," 11.25 mm: Comb on apex of
article 6 on pereopod 3 absent, article 2 of pereopod
5 with only 6 facial setae; inner ramus of uropod 1 so
cylindrical that dorsal (medial) spines rotated out of
sight medially (see illustration), peduncle of uropod
2 with 6 dorsal spines, outer ramus with 8, inner
with 4.
DESCRIPTION OF MALE.?Eyes enlarged. Article 1 of
antenna 1 with increased numbers of ventral setules,
no medial fuzz; primary flagellum with 9 articles,
one calceolus each on articles 1-4, aesthetascs well
developed; accessory flagellum with 8 articles. Facial
spine formula on article 4 of antenna 2 = 2-12-11-7,
dorsomedial margin fuzzy, other setae shortened;
article 5 with 8 dorsal sets of male setae and 7 cal-
ceoli, ventrodistal apex with 2 setules; flagellar for-
mula = 45, 2-4, 6, 8 . . . 40 or 44, 2-5, 7, 9 . . . 39.
Distal member of right lacinia mobilis poorly sub-
bifid, inner margin of article 3 on mandibular palp
with 2 accessory setae not in female. Inner plate of
maxilliped with 2 main apical spines spread slightly
more than in female, 4 medial setae; outer plate
with 12 spines; palp article 2 with 3 lateral setae,
article 3 with 6 lateral setae. Coxae 3-4 slightly to
much longer than in female; ventral setal formula
of coxae 1-4 = 10-13-13-27. Long posterior setae
on article 2 of gnathopods 1-2 and pereopods 1-2 =
6-6-6-3, long anteriors = 6-9-14-7, short anteri-
ors = 1-2-4-3, no others. Setal and spine formulas
of pereopods 1-2 on article 4 = 5 + 5 (separated by
gap from other distal setae), on article 5 = 5 and 5,
on article 6 = 9 + 10 plus middistal spine, setae on
posterodistal margin of article 4 shorter than in
female. Article 2 of pereopods 3 and 5 slightly nar-
rower than in female. Epimera 1-2 with numerous
short posterodorsal setae, epimera 1-3 slightly
broadened, posterior margin of epimeron 3 not
shortened. Urosome ventrally with 3 paired sets of
setae, lateral set = 7, ventral sets = 3 and 4. Spine
formula of uropods: uropod 1 peduncle apico-
lateral = 3; uropod 2 peduncle dorsal = 6; dorsal
spines on outer ramus of uropod 1 = 9 , continuous
to apex, but with slightly excessive gap between
spines 1-2, of uropod 2 = 8; inner ramus of uropod
1 = 6-7 plus 3, of uropod 2 = 3. Ventral spines on
peduncle of uropod 3 = 14-15; spine formula on
article 1 of outer ramus = 0-1-1-2-2-2 or 0?1?1?1?
2-2-2-2, setal formula = 3-1-1-1-1-1 or 2-1-2-1-
1-1-1-1. Telson weakly elongate, broadened api-
cally, distal spines scarcely shortened, midbase of
each lobe with numerous acute denticles.
VARIATIONS.?Aberrant male, 8.0 mm, PPBES
960/3: Eyes scarcely enlarged but penial processes
well developed; antenna 2 like that of female; uro-
pod 3 almost fully masculine but antenna 1 and
epimera like those of female.
ILLUSTRATIONS.?Female maxilla 2 as in Tipi-
megus thalerus; several spines on ventral side of
peduncle on uropod 3 omitted.
HOLOTYPE.?AM, female "n," 7.60 mm.
TYPE-LOCALITY.?SBS C4S4, 17 May 1972, off
Malabar, New South Wales, Australia, 69 m, sand.
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
VOUCHER MATERIAL.?SBS E2S4: female "w," 7.20
mm (illus.); male "y," 7.10 mm (illus.); male "u,"
6.72 mm (illus.); juvenile "j ," 4.37 mm (illus). CPBS
24N: female "z," 11.25 mm (illus.).
RELATIONSHIP.?This species differs from Tipi-
megus thalerus in the elongate dactyls of pereopods
3-4, the small size of the cuticular setules, the broad
apices of the hands on the gnathopods, the widely
spread setae on coxae 1-3, and the elongation of the
lateral seta on article 2 of uropod 3 which in T.
thalerus forms a shorter and distinctive spine. Tipi-
megus kangulun is characterized by the noncontinu-
ous dorsal spination on uropod 1, particularly well
developed in females and juveniles but becoming
obscured in males. Male uropod 1 usually has some
weak degree of asymmetry in the placement of
apical spines and lacks any definite apical spine
fully contiguous with the nail. In all but super-
females article 6 of pereopod 3 bears a distal comb
which is absent in T. thalerus. Generally the spine
forming part of the spike on the peduncle of uro-
pod 1 is shorter in T. kangulun than in T. thalerus.
REMARKS.?A single male with stunted antenna 2
from JLB AUS 11, Western Australia, is provision-
ally identified with T. kangulun but the setae on
coxae 1-3 are few and narrowly spread; this may be
a new species and will be reserved until more mate-
rial from Western Australia can be collected.
MATERIAL.?CPBS, 3 samples from 2 stations (3);
WPBES, 2 samples from 2 stations <5); PPBES, 14
samples from 10 stations (26) [5 doubtful from 2
samples]; AM, one sample (1); SBS, 6 samples from
4 stations (17).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 2-13 m, sand. New South Wales: off
Sydney, 53-150 m, sand.
Tipimegus kalkro, new species
FIGURES 20-23
DESCRIPTION OF FEMALE.?Head about 22 percent
of total body length, greatest width about 70 per-
cent of length, rostrum constricted, narrow, reach-
ing apex of article 2 on antenna 1. Eyes medium,
partly occluded with pigment. Article 1 on peduncle
of antenna 1 about 1.2 times as long as wide, about
twice as wide as article 2, ventral margin with about
11 setules, weakly produced dorsal apex with 2
setules; article 2 about 1.1 times as long as article
1, with ventral crescent of 12-13 setae; primary fla-
gellum with 8-9 articles, about 0.65 times as long as
peduncle, bearing long aesthetascs; accessory flagel-
lum with 8 articles. Spine formula on article 4 of
antenna 2 = 11-8-5 or 11-8-4 + 3-4 supernumer-
ary spines, dorsal margin setose, ventral margin
with 7-8 groups of 2-5 long to medium setae, dense
ventrodistal brush of setae; article 5 about 0.45 times
as long as article 4, facial spine formula = 0, dorsal
margin sparsely setulose. ventral margin with 9
unpaired setae, 5 ventrodistal medium spines, this
row set facially: flagelluin about 0.9 times as long as
articles 4-5 of peduncle combined, with 11-12 arti-
cles. Mandibles with weak palpar hump: right in-
cisor with 4 teeth; left incisor with 4 teeth in 2
humps; right lacinia mobilis bifid, distal branch
much shorter than proximal, broad, subhind, proxi-
mal branch simple, pointed; left lacinia mobilis
with 4 teeth; right rakers 10-12 plus 0-1 rudimen-
tary; left rakers 13-14 plus one rudimentary; molars
elongate, conical or columnar, each with one long
apical spine, one short apical spine, one medium
spine on side with vestigial nonarticulate partner;
palp article 1 short, article 2 with 1-2 medium in-
ner apical setae, plus 3-5 other shorter apical setae
in clump and one other short inner seta, and 3
outer facial setae, article 3 about 0.95 times as long
as article 2, oblique apex with 16 spine-setae, baso-
dorsal formula = 3 (or 2 on left occasionally). Inner
plate of maxilla 1 large, bearing one long subapical
pluseta, 2 apicolateral shorter setae; palp article 2
with 7-8 apical spines and 12 submarginal and facial
setae. Inner plate of maxilla 2 shorter and narrower
than outer, outer lacking apicolateral setae, inner
with 6 medial setae. Inner plate of maxilliped with
2 large, thick apical spines, 6 apicofacial setae, 4
medial setae, outer plate with 14 medial and apical
spines, no apicolateral setae; palp article 1 lacking
apicolateral seta, article 2 with one group of 2
apicolateral setae, medial margin of article 2
strongly setose, article 3 with 29 facial setae, 5
lateral setae, article 4 with 3 apical setules, nail
absent. Coxa 1 strongly expanded distally, anterior
margin straight; main ventral setae of coxae 1-4 =
(6-7H9-H)- (9 - l lH16-22) , posteriormost seta of
coxae 1-3 shortest; anterior and posterior margins
of coxa 4 strongly divergent, posterior margin
oblique, almost straight, posterodorsal corner sharp,
posterodorsal margin ordinary, almost straight;
width-length ratio of coxa 4 = 17:18. Long posterior
NUMBER 245 65
FICURE 20.?Tipimegus kalkro, new species, holotype, female "d," 5.66 mm (b ? male "b," 5.00
ram; c ? female "c," 5.43 mm; k = female "k," 5.67 mm).
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 21.?Tipimegus kalkro, new species, holotype, female "d," 5.66 mm (fc ? male "b," 5.00
mm; c ? female "c," 5.43 mm).
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 6-8-9-7, long anteriors = 17(+ 3 facial)-14-
15-1, short anteriors = 2-3-4-4, no others. Gna-
thopods with thin hands narrow apically; width
ratios of articles 5-6 on gnathopods 1-2 = 30:21
and 29:21, length ratios = 65:50 and 63:47, palmar
humps small, palms transverse; article 5 of gnatho-
pods 1-2 elongate, ovate, posterior margins rounded.
Pereopod 2 stouter than pereopod 1, especially arti-
cle 4; facial setae formula on article 4 = 7 or 8 and
8 or 8, on article 5 = 6 and 6 or 6 and 7; main
spine of article 5 extending to M. 85 on article 6,
spine formula of article 6 = 9 + 9 and 9 + 9 or
8 + 9 plus huge middistal spine; acclivity on inner
margin of dactyls of pereopods 1-2 sharp, produced
as tooth, emergent setule short, midposterior pluseta
short. Coxae 5-7 posteroventral seta formula = 22-
35-30 composed of marginal and facial groups. Arti-
cles 4-5 of pereopods 3-4 broad, facial spine rows
dense; facial ridge formula on article 2 of pereopods
3-5 = 0-1-1; width ratios of articles 2, 4, 5, 6 of
pereopod 3 = 53:47:48:22, of pereopod 4 = 57:55:
43:19, of pereopod 5 = 55:9:8:4; length ratios of
pereopod 3 = 72:42:41:31, of pereopod 4 = 64:57:
NUMBER 245 67
bP2
FIGURE 22.?Tipimegus kalkro, new species, holotype, female "d," 5.66 mm (b ? male "b," 5.00 mm).
42:40, of pereopod 5 = 75:20:19:20; article 2 of
pereopod 5 reaching middle of article 5, ventrally
setose; medial apex on article 6 of pereopods 3 and
5 finely combed; dactyls of pereopods 3-4 short.
Posteroventral corner of epimeron 1 weakly protru-
sive, posterior margin weakly convex, setose, antero-
ventral face with 7-8 short to medium setae, mid-
face with 11 medium setae; posteroventral corner of
epimeron 2 rounded, weakly protuberant, posterior
margin convex, setose, facial setae = 8-9 plus one
midfacial, some pairs set vertically (rows of 7 + 2 or
6 + 2); posteroventral corner of epimeron 3 weakly
protuberant, posterior margin almost straight,
weakly serrate, setose, ventral margin with 8?10
setae mainly in posterior half, face with several sub-
marginal setae. Urosomite 1 with 4 paired sets of
setae, each side with sets of 5-7, ventrally with sets
of 2-A, 2-4, and 2, articulation line complete; uroso-
mite 3 protuberant dorsally. Rami of uropods 1-2
with articulate enlarged sharp apical nails, rami of
uropods 1-2 with 1-2 accessory nails, continuously
spinose; outer ramus of uropod 1 with 7 dorsal
68
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
\
bT
FIGURE 23.?Tipimegus kalkro, new species, holotype, female "d," 5.66 mm (b ? male "b," 5.00
mm; c ? female "c," 5.43 mm; y ? male "y," 5.30 mm).
NUMBER 245 69
spines, inner with 5-8 dorsal plus 3-5 lateral; outer
ramus of urojxxl 2 with 4-5 dorsal spines, inner
with 5 dorsal spines; peduncle of uropod 1 with 2
apicolateral spines, medially with 1-2 marginal
spines; peduncle of uropod 2 with 3 dorsal spines,
medially with one small apical spine; outer rami
of uropods 1-2 and apex of peduncle on uropod 2
with large comb (or comb absent on peduncle of
uropod 2). Peduncle of uropod 3 with 9-12 ventral
spines, dorsally with one lateral spine, medial spine
and setule or 2 spines only; rami submasculine,
inner extending to M. 100+ on article 1 of outer
ramus, apex with 3 setae, medial margin setose,
lateral margin with one apical seta; article 2 of
outer ramus ordinary, 0.20, bearing 3 medium to
long setae, lateralmost shortest, thin, nonplumose,
spiuclikc; apicomedial margin of article 1 with 3
setae, lateral margin with 4 acclivities, spine form-
ula = 2-2-2-2, setal formula = 0. Telson ordinary,
length-width ratio = 1:1, not fully cleft, each apex
of medium width, rounded, undulant, lateral accliv-
ity shallow, weak, with lateral and medial spines
separated by setule, midlateral setules diverse. Cuti-
cle with ordinary bulbar setules, surface with dense
fuzz of denticles, emergent setules branched.
OBSERVATIONS (female).?Article 2 on pereopod 5
of female "c" with 11 long facial setae, 9 setules but
only 3 anterodorsal setae; article 5 of pereopod 1
in adults with strong medial comb on distal bevel,
with one medioproximal spinule; female "d" holo-
type with aberrant right lacinia mobilis, proximal
branch stubby, perhaps broken and regenerate.
DESCRIPTION OF MALE.?Less robust than female.
Eyes enlarged. Article 1 of antenna 1 lacking medial
fuzz, bearing 16 or more ventral setules; article 2
with 11 ventral setae; one calceolus each on articles
1-3 of primary flagellum. Spine formula on article 3
of antenna 2 = 3 plus disjunct setule, on article
4 = 3-9-7-6 or 3-11-7-5, (or, rarely, variations
thereof), plus 3 supernumerary spines, on article
5 = 2, article 4 with dorsomedial fuzz, ventral setae
shortened and fewer than in female; article 5 with
5 distoventral setules, dorsal margin with 5-6
calceoli and 7-8 sets of male setae; flagellum with
approximately 42 articles (often broken), flagellar
formula = 42, 3, 4, 6, 8 . . . 40 (calceoli on segments
3-12 rudimentary), or 2-7, 9 . . . (calceoli on 5, 6, 7,
11 rudimentary), or 2-5, 7, 9 . . . (calceoli on 3 and
7 rudimentary), or, rarely, other variations. Upper
lip more deeply incised than in female. Mandibular
molars bearing only 3 spines; right rakers 10 plus
one rudimentary, left 13 plus one rudimentary;
basofacial setal formula on palp article 3 = 2, inner
margin with 5 setae (absent in female). Coxa 4
narrower than in female but of similar length rela-
tive to coxa 1. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 1-4-5-4, long
anteriors = 10-11-12-1, short anteriors = 1-3-5-3.
Pereopods 1-2 thinner than in female; facial and
spine formulas on article 4 = 7 and 6, on article
5 = 5 and 5, on article 6 = 8 + 8 plus huge mid-
distal spine. Article 2 of pereopod 5 narrower
than in female; articles 5 and 6 of pereopod 4
greatly to slightly narrower. Epimera 1-3 broad-
ened; posterior margin of epimeron 3 shortened;
facial ridge on epimeron 2 weak or absent; setal
formulas, epimeron 1 anteroventral = 5-6, facial =
14 decreasing to 5, epimeron 2 facial = 5-6, epi-
meron 3 ventral = 8-9. Spine formulas of uro-
pods, uropod 1 peduncle apicolateral = 2, me-
dial = 1, uropod 2 peduncle dorsal = 6-7, dor-
sal spines on outer ramus of uropod 1 = 8, of
uropod 2 =6-7, inner ramus of uropod 1 = 5 + 5,
of uropod 2 = 3. Ventral spines on peduncle of
uropod 3 = 12-13, spine formula on article 1 of
outer ramus = 0-0-0-1-1-2-2-3-3, or 0-0-0-2-2-2,
or, rarely, other variations thereof, setal formula =
1 X 9 , or 2-1-1-1-1-0-0, or other variations.
Telson weakly elongate, broadened, 2-3 distal spines
on each lobe shortened, each lobe with patch of
midbasal denticles.
ILLUSTRATIONS.?Lateral head of female "c" illus-
trated but outline of antenna 1 added to drawing
from holotype female "d" by adjusting heads to
same proportions; palp of maxilla 1 heavily flat-
tened, 2 setae omitted; palp article 3 of maxilliped
with several setae omitted from face on whole view
but enlargement with setae marked by their pits
showing all 29 facial setae; article 2 on outer ramus
of male uropod 3 lacking mid seta, often broken
off, lateralmost spine-seta plumose or not (variable
on single individual); denticles of male telson too
small to show on illustration.
HOLOTYPE.?NMV, female "d," 5.66 mm.
TYPE-LOCALITY.?PPBES 912/4, 19 Nov 1971,
Port Phillip Bay, Victoria, Australia, 7 m, sand.
VOUCHER MATERIAL.?Type-locality: female "c,"
5.43 mm (illus.); young male "a," 5.72 mm. PPBES
912/1: male "b," 5.00 mm {illus.); male "y," 5.30
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
mm (illus.). PPBES 920/5: female '%" 5.67 mm
(illus.); male "m," 5.61 mm.
RELATIONSHIP.?This species differs from T.
thalerus, T. dinjerrus, and T. kangulun in the
presence of grossly enlarged combs on the outer
rami of uropods 1-2. It further differs from T.
thalerus in the smallness of the cuticular setules,
in the greater elongation or thinness of the special
lateral spine-seta on article 2 of the outer ramus
on uropod 3, in the fewer spines on the outer
ramus of uropod 2, the greater number of spines
on the inner ramus of uropod 2, the fewer spines
medially on the peduncle of uropod 1, and in the
longer spines on uropods 1-2. From T. dinjerrus,
T. kalkro further differs in the narrow apices of
the gnathopods. From T. kangulun, T. kalkro fur-
ther differs in the evenly spread spination on the
outer ramus of uropod 1, in the shortened dactyls
of pereopods 3-4, and in the narrow apices of the
gnathopods.
MATERIAL.?PPBES, 23 samples from 13 stations
(54).
DISTRIBUTION.?Victoria, Port Phillip Bay, 4-8
m, sand.
Tipimegus stebbingi (J. L. Barnard),
new combination
Pontharpinia pinguis.?Stebbing, 1897:35-34, pi. 92J [not Has-
well, 1880]; 1910:635
Paraphoxus stebbingi J. L. Barnard, 1958:148; 1960:282.
TYPE.?Female 3/10 inch.
TYPE-LOCALITY.?Jervis Bay, New South Wales,
Australia.
NOTES.?The female identified by Stebbing as
Pontharpinia pinguis (Haswell) does not belong to
that species, nor, indeed to the genus Pontharpinia.
Stebbing used this female to erect the genus
Pontharpinia to contain Pontharpinia pinguis as
the type-species in monotypy. Pontharpinia is there-
fore a confounded genus but we prefer to maintain
Pontharpinia as a valid genus for the purposes of
nomenclatural stability. J. L. Barnard (1958) re-
named Stebbing's specimen but according to Dr.
Roger J. Lincoln of the British Museum (Natural
History), Dr. D.J.G. Griffin of the Australian
Museum and Dr. Torben Wolff of the Universitetets
Zoologiske Museum, this specimen is lost. Those
are the three museums in which it might have been
deposited.
We cannot identify T. stebbingi from Stebbing's
description; extensive exploration of Jervis Bay
will be necessary to sort out the probable identity.
Tipimegus stebbingi is apparently not in the genus
Booranus because article 2 of pereopod 5 is poorly
inflated anteriorly in that genus. The short dactyl
of pereopod 3 and the apically narrowed hands of
the gnathopods suggest that T. stebbingi may be
identical to T. thalerus or T. kalkro but the widely
spread setae of coxa 2 suggest affinity with T.
kangulun. Tipimegus stebbingi may therefore be
fully distinct from any of the new species described
herein.
Tipimegus species 4N
NOTES.?This probable new species is represented
in the collections only by an immature male; its
full description is reserved until additional mate-
rials become available. Setae on coxa 1 poorly
spread; hands of gnathopods broad apically, palms
straight; dactyl of pereopod 3 short, article 6 stout,
dactyl of pereopod 4 longer than half of article 6,
latter elongate, thin; outer ramus of uropod
1 with continuous and apically dense dorsal spina-
tion; outer ramus of uropod 2 with dense and even
but fine dorsal comb; epimeron 2 bearing setae in
middle of lateral surface; cuticular setules com-
posed of very small bulbs, emergent setule at most
asymmetrically bifid or bearing very few side
plumes.
It differs from T. thalerus in the very elongate
article 6 and dactyl of pereopod 4, the presence of
midsetae on epimeron 2, and the dense clumping
of apical spines on the outer ramus of uropod 1.
It also differs from T. dinjerrus in the presence
of midsetae on epimeron 2, the shorter dactyl of
pereopod 4 and the dense clumping of apical spines
on the outer ramus of uropod 1.
MATERIAL.?SBS E253, male, 3.84 mm, New
South Wales, off Malabar, 52 m, sand.
Tipimegus species 5N
NOTES.?This apparently distinct species is repre-
sented in the collections by subadult males and
juveniles; its full description is reserved until adult
females become available. Setae on coxa 1 poorly
spread; hands of gnathopods narrow apically (but
slightly broader than in T. thalerus), palm of
gnathopod 1 S-shaped; dactyls of pereopods 3-4
longer than in T. thalerus but less than half as
NUMBER 245 71
long as article 6, latter elongate and thin in both
pereopods 3-4; outer ramus of uropod 1 with con-
tinuous spination but with weak and excessive gap
between spines 1 and 2 (spine 1 closely adjacent to
nail); outer ramus of uropod 2 lacking conspicu-
ous dorsal comb but with ultrafine ragged comb-
fuzz; epimeron 2 lacking setae in middle of lateral
surface; cuticular setules composed of small bulbs
of size similar to those of T. dinjerrus, emergent
setules branched.
It differs from T. thalerus in the weakly S-shaped
curve on the palm of gnathopod 1, in the greater
elongation of the dactyl and article 6 of pereopods
3-4, and in the greater gap between spines 1 and
2 on the outer ramus of uropod 1.
It differs from T. species 4N in the S-shaped but
slightly shorter palm of gnathopod I, the absence
of middle setae on epimeron 2, the sparser apical
spination on the outer ramus of uropod 1, and the
somewhat shorter dactyls of pereopods 3-4.
It further differs from T. dinjerrus in the S-shaped
palm of gnathopod 1 and the slightly more elon-
gate articles 6-7 of pereopods 3-4.
MATERIAL.?PPBES, 7 samples from 3 stations
(18).
DISTRIBUTION.?Victoria, Port Phillip Bay, 4-8
m, sand.
Boor anus, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
ordinary, ventral setae widely spread. Article 1 of
antenna 2 weakly ensiform; article 3 with num-
erous setae, setules and spines; facial spines on arti-
cle 4 in 2 or more rows, plus special apical spines;
article 5 especially thin and short. Right mandi-
bular incisor with 4 teeth; molar not triturative,
large, elongate conical, then subtruncate and bear-
ing 3-4 special large spines, not bearing fuzz; palpar
hump small. Palp of maxilla 1 biarticulate; inner
plate with 4-5 setae. Setation of maxilla 2 ordinary.
Inner plate of maxillipeds ordinary; apex of palp
article 3 not protuberant, dactyl elongate, apical
nail absent. Gnathopods small, similar; article 5 of
gnathopods 1-2 elongate, free, without eusirid at-
tachment; palms transverse to chelate, hands elon-
gate, heavily setose anteriorly, trichophoxid in
shape. Article 5 of pereopods 1-2 lacking setae
posteroproximally. Article 2 of pereopod 3 of broad
form, articles 4-5 of pereopods 3-4 broad, article 2
of pereopods 3-4 setose posteriorly; pereopod 5
especially small, article 2 strongly setose ventrally,
article 3 greatly enlarged, dactyl normal, pereopod
5 with gills. Epimera 1-2 bearing numerous long
posterior setae, rarely with midfacial setae above
ventral facial ridge; epimeron 3 ordinary, bearing
row of facial spines. Urosomite 1 naked laterally,
bearing one or more midventral crescents or bun-
dles of setae; urosomite 3 lacking dorsal hook or
special process. Peduncle of uropod 1 normally
elongate to weakly shortened, with apicoventral
spike, without special enlarged apicolateral or
medial spine, only pedunclar apices of uropod 2
combed; inner ramus of uropod 1 with 2 rows of
marginal spines, some rami continuously spinose to
apex; inner ramus of uropod 2 especially short-
ened. Uropod 3 ordinary, article 2 of outer ramus
carrying 3 short to long apical setae. Telson
ordinary, with 2-3 apical spines or setae on each
lobe plus setules.
DESCRIPTION.?Rostrum fully developed, con-
stricted. Fuzz on article 1 of antenna 1 in male
absent; calceoli on male primary flagellum of
antenna 1 present. Calceoli on article 5 of male
antenna 2 present, facial spines absent in female,
flagellum in male with calceoli. Prebuccal parts
strongly extended forward, especially massive,
strongly distinct, epistome with large tooth, neither
epistome nor upper lip dominant. Right lacinia
mobilis bifid, thin; article 1 of mandibular palp
short, palp medium, apex of article 3 oblique, arti-
cle 2 with outer setae. Lower lip lacking cones.
Outer plate of maxilla 1 with 11 spines, one spine
especially thickened. Inner plates of maxilliped
thick, ordinarily setose. Coxae 2-4 with special
anterodorsal humps. All posterior spines on article
6 of pereopods 1-2 thick and stiff, midapical spine
present, especially enlarged. Article 2 of pereopod
5 with facial setae. Peduncle of uropod 1 with
dorsolateral spines widely spread, medial spines
partially spread; peduncle of uropod 2 with medial
spines confined apically. Peduncle of uropod 3 lack-
ing extra subapical setae or spines. Telson with
ordinary pair of midlateral or dorsal setules on
each side.
TYPE-SPECIES.?Booranus weemus, new species.
COMPOSITION.?Booranus tikeri, new species; B.
wangoorus, new species.
RELATIONSHIP.?This genus is extremely close to
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Tipimegus but is believed to qualify as a full and
separate genus because each forms a tight cluster of
3 to 5 species without intergradations. The main
distinctions of Booranus are the massively produced
epistome and the presence of gills on pereopod 5.
The two clusters also differ in the presence (Boor-
anus) or absence (Tipimegus) of a grossly devel-
oped lateral spine row on epimeron 3.
See "Relationship" under Tipimegus for other
common characteristics.
Key to the Species of Booranus
1. Article 5 of pereopod 5 lacking apical comb laterally B. tikeri, new species
Article 5 of pereopod 5 bearing apical comb laterally 2
2. Coxa 7 with 1-2 facial setae (apart from posterior setae), spike on uropod I reaching to M. 50
on outer ramus, epimeron 1 with a few midfacial setae B. wangoorus, new species
Coxa 7 lacking facial setae, spike on uropod 1 reaching to M. 30-35 on outer ramus, epimeron
1 lacking facial setae in middle above lateral ridge and apart from posterior setae
B. iveetmu, new specie*
Booranus weemus, new species
FIGURES 24-26
DESCRIPTION OF FEMALE.?Head about 22 percent
of total body length, greatest width about 60 per-
cent of length, rostrum constricted, narrow, elon-
gate, exceeding middle of article 2 on antenna 1.
Eyes large, clear of pigment, ommatidia ordinary.
Article 1 on peduncle of antenna 1 about 1.25
times as long as wide, about twice as wide as article
2, ventral margin with about 4 setules, weakly pro-
duced dorsal apex with 1-3 setules; article 2 about
as long as article 1, with ventral cycle of 12-13
setae; primary flagellum with 10-11 articles, about
0.85 times as long as peduncle, bearing medium
aesthetascs; accessory flagellum with 11 articles.
Article 3 of antenna 2 with 5 setae-spines and one
setule; spine formula of article 4 = 4-10-9-9 or
3-9-10-8, dorsal margin with 4 notches each bear-
ing 3-5 setae, ventral margin with 7 groups of
1-5 long to short setae, 2-3 distal long spines and
brush of 11 + setae; article 5 about half as long as
article 4, facial spine formula = 0, dorsal margin
bearing 2 setae, ventral margin with 7-9 unpaired
setae, clump of 5-6 ventrodistal long to short
spines, no subdistal facial spines; flagellum about
as long as articles 4-5 of peduncle combined, with
14 articles, epistome with sharp anterior protru-
sion; upper lip with ventral sinus. Mandibles with
weak palpar hump; right incisor with 4 teeth; left
incisor with 4 humps in 2 branches; right lacinia
mobilis bifid, distal branch shorter than proximal,
subbifid, proximal branch simple, pointed, with
marginal denticles, humps; left lacinia mobilis with
4 teeth; right rakers 11 plus or minus one rudi-
mentary; left rakers 12-13; molars elongate, coni-
cal, each molar with 4 long to short spines, none
disjunct; palp article 1 short, article 2 with one
long and one short inner apical setae and 6 other
shorter inner and facial setae, article 3 about 1.1
times as long as article 2, oblique apex with 12
spine-setae, basodorsal formula = 0-2. Inner plate
of maxilla 1 large, bearing one long apicofacial
pluseta, 3 shorter apical setae; palp article 2 with
9-10 apical marginal spines and 9-10 submarginal
setae. Inner plate of maxilla 2 shorter and narrower
than outer, outer lacking apicolateral setae, inner
with 10-11 medial setae. Inner plate of maxilliped
with 2 large thick apical spines, 5-6 apicofacial
setae, 4-5 medial setae; outer plate with 15-18
medial and apical spines, one apicolateral seta;
palp articles 1-2 lacking apicolateral setae; medial
margin of article 2 moderately setose; article 3 with
about 30 facial setae, 7-9 lateral setae in 5 groups;
article 4 with 5 accessory setules, mostly on face,
nail absent. Coxa 1 strongly expanded distally, an-
terior margin almost straight; main ventral setae
of coxae 1-4 = (!^>)-(?-Wy-(9-\0)-\l, posterior-
most seta of coxae 1-3 shortest; anterior and poste-
rior margins of coxa 4 divergent, posterior margin
oblique, almost straight, posterodorsal corner
rounded, posterodorsal margin short, undulant,
width-length ratio of coxa 4 = 13:16; gills present
on coxae 2-7. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 4-3-7-6, long
anteriors = 20(+ 3 facial)-7-8-(2-3), short anteri-
ors = 3-5-4-(4-5), no others. Gnathopods thin,
hands thin, parachelate, mitellid (occasionally
NUMBER 245 73
FIGURE 24.?Booranus weemus, new species, female "a," 6.1 mm (g ? holotype female "g," 5.63
mm; n ? male "n," 5.58 mm).
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Gl
FIGURE 25.?Booramts weemus, new species, female "a," 6.1 mm (Y m dactyl).
gnathopod 2 slightly smaller than gnathopod 1);
width ratios of articles 5-6 on gnathopods 1-2 =
29:24 and 28:25, length ratios = 63:46 and 65:46;
palmar humps very small, palms transverse; article
5 of gnathopods 1-2 elongate, ovate, posterior mar-
gins rounded-flat. Pereopods 1-2 similar; article 4
of pereopods 1-2 similar but much stouter on
pereopod 2; article 5 with apicomedial comb on
pereopods 1-2; facial setae formula on article 4 =
5 and 5 or 5 and 4, on article 5 = 5 and 6 or 6 and
6; main spine of article 5 extending to M. 75 on
article 6, article 5 with one extra long distopos-
terior spine and numerous setae, proximal margin
naked; spine formula of article 6 = 6 + 8 and 7 + 8
or 7 + 9 and 8 + 9 plus huge middistal spine, some
spines especially long; acclivity on inner margin of
dactyls of pereopods 1-2 sharp, produced as tooth,
emergent setule short, marginal pluseta highly proxi-
mal. Coxae 5-7 posteroventral seta formula = 18-
18-11, none facial. Articles 4-5 of pereopods 3-4
broad, facial spine rows dense; facial ridge formula
on article 2 of pereopods 3-5 = 0-1-1; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 52:48:44:22, of
pereopod 4 = 65:55:34:14, of pereopod 5 = 60:10:
8:4; length ratios of pereopod 3 = 71:39:38:35, of
pereopod 4 = 77:59:51:52, of pereopod 5 = 89:20:
21:22; article 2 of pereopod 5 almost reaching apex
of article 5, ventrally setose, anterior margin weakly
NUMBER 245 75
w
FIGURE 26.?Booranus weemus, new species, female "a," 6.1 mm (g
mm; n -= male "n," 5.58 mm).
holotype female "g," 5.63
bulging, pereopod 5 small and mostly hidden by
pereopod 4, lateral apex of article 5 and medial
apex of article 6 finely combed. Posteroventral
corner of epimeron 1 rounded-quadrate, posterior
margin straight, weakly setose (4), anteroventral
margin and face with 6-7 short setae, posteroventral
margin with 2-3 short setae, no fully facial setae;
posteroventral corner of epimeron 2 rounded, post-
erior margin straight, weakly serrate, setose, facial
setae = 5-7, none set vertically; posteroventral cor-
ner of epimeron 3 rounded, weakly protuberant,
posterior margin almost straight, serrate, setose,
ventral margin with 6-7 setae mainly in posterior
half, face with oblique row of 7-8 spines in mid-
dle; epimera 1-2 with setule on posterodorsal mar-
gin set in weak notch. Rami of uropods 1-2 with
articulate enlarged sharp apical nails and accessory
nails, spike on uropod 1 reaching to M. 30-35 on
outer ramus, outer ramus of uropod 1 with 8-10
dorsal spines, inner with 5-6 and facial row of %-A
spines; outer ramus of uropod 2 with 5-6 dorsal
spines, inner with 3-4 dorsomedial spines includ-
ing accessory nail; peduncle of uropod 1 with 4
apicolateral spines, no basofacial setae, medially
with 3 marginal spines, apicalmost ordinary; ped-
uncle of uropod 2 with 4?5 dorsal spines, medially
with pair of apical spines; apicolateral corners of
peduncles on uropods 1-2 with weak comb. Ped-
uncle of uropod 3 with 11-12 ventral spines, dor-
sally with 4-5 lateral spines, one medial spine;
rami masculine, inner extending to M. 100 on
article 1 of outer ramus, apex with 3 setae, medial
and lateral margins setose; article 2 of outer ramus
ordinary, 0.17, bearing 2 medium setae and one
spine, apicomedial margin of article 1 setose, lat-
eral margin with 7-8 acclivities, spine formula =
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(l)_2-2-2-2-2-2-2-2, one spine of each set elon-
gate, setal formula = 0. Urosomite 1 with midlateral
group of 8-9 setae, one ventral seta on each side,
articulation line incomplete; urosomite 3 protu-
berant dorsally, with middorsal spine on each side.
Telson long, length-width ratio = 31:28, not fully
cleft, each apex narrow to medium, rounded, with
middle acclivity, lateral acclivity narrow, weak,
bearing ordinary lateral spine, spine next medial
longer, with lateral and medial spines separated by
setule, often with third spine fully lateral, mid-
lateral setules diverse. Cuticle with bulbar setules
closely packed with pipes and spicules or studs,
setules surrounded by cordate plaques, surface
bearing fine striations in form of linear fingerprint
pattern, emergent setules branched.
OBSERVATIONS (female).?Description based on
holotype female "g" and female "a." Female "a"
with aberrant, narrowed telson bearing only 2
(normal) apical spines on each lobe; spike of uropod
1 about 15 percent shorter than in female "g";
facial ridges on article 2 of pereopod 3 probably
artifacts.
DESCRIPTION OF MALE.?Eyes hugely enlarged.
Article 1 of antenna 1 lacking medial fuzz, bearing
numerous ventral setules; article 2 with 11 ventral
setae; primary flagellum with 11 articles, one cal-
ceolus each on articles 1-5, aesthetascs on primary
flagellum as in female; accessory flagellum with 9
articles. Facial spine formula on article 4 of antenna
2 = 4-10-9-7; article 5 with 5 ventral setae, one
subapical facial spine, one setule, proximal face
with one setule, dorsal margin with 4 sets of male
setae and 3 calceoli, ventrodistal apex with 5 setules
in row; flagellar formula = 28, 2, 4, 6 . . . 14. Man-
dibular palp article 3 as in female, lacking setae on
inner margin. Coxa 4 similar to but smaller in rela-
tion to coxa 1 than in female; ventral setal formula
of coxae 1^1 = 8-9-9-15. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
2-3-3-3. Pereopods 1-2 much thinner than in fe-
male; facial and setal spine formulas of pereopods
1-2 on article 4 = 4 and 4, on article 5 = 5 and 6,
on article 6 = 6 + 8 and 7 + 8 plus middistal spine.
Article 2 of pereopods 3, 4, 5 slightly narrower than
in female. Epimera 1-3 broadened; posterior mar-
gin of epimeron 3 not shortened; setal formulas,
epimeron 1 anteroventral = 4, ventral = 7 widely
spread, posterior = 8, epimeron 2 facial = 5, poste-
rior = 8, epimeron 3 posterior = 8, facial = 9, ven-
tral = 5. Spine formulas of uropods, uropod 1 pe-
duncle apicolateral = 4, uropod 2 peduncle dorsal =
6, dorsal spines on outer ramus of uropod 1 = 10
including nails, of uropod 2 = 9 including nails, in-
ner ramus of uropod 1 = 6 and 5 including nails, of
uropod 2 = 5 including nails, spine formula on arti-
cle 1 of outer ramus = 0-0-0-2 X 8, spines shorter
than in female, setal formula = 1 x 1 1 . Telson
broadened, distal spines shortened, each lobe with
broad patches of dorsal denticles.
OBSERVATIONS (male).?Male "n" dimensions of
pereopods 3-5, widths of articles 2, 4, 5, 6 of pereo-
pod 3 = 50:44:42:20, of pereopod 4 = 60:50:30:13.
of pereopod 5 = 54:10:8:5, lengths of articles on
pereopod 3 = 77=40:35:36. of pereopod 4 = 62:65:
50:52, of pereopod 5 = 90:21:23:23; alternative spine
formula on outer ramus of uropod 3 = 0-1-2-2-2
. . . n.
ILLUSTRATIONS.?Generally as in Booranus tikeri
except in specific characters mentioned for female
or for notes in male description. Following female
parts like Tipimegus thalerus except as specified in
description: maxilla 2 and maxilliped. Following
parts like B. tikeri except as specified in description:
lower lip and maxilla 1 except for more elongate
inner plate as figured for B. tveemus.
HOLOTYPE.?NMV, female "g," 5.63 mm.
TYPE LOCALITY.?CPBS 12S/1, 16 Mar 1965,
Western Port, Victoria, Australia, 2.4 m, fine sand,
mud.
VOUCHER MATERIAL.?Type-locality: female "a,"
6.1 mm (illus.); male "n," 5.58 mm (illus.); smallest
juvenile examined, 3.5 mm.
MATERIAL.?CPBS, 5 samples from one station
(38); WPBES, one sample <2).
DISTRIBUTION.?Victoria, Western Port, 0.3-2.0 m,
sand, sand with mud, seagrass.
Booranus tikeri, new species
FIGURES 27, 28
DESCRIPTION OF FEMALE.?Head about 20 percent
of total body length, greatest width about 65 per-
cent of length, rostrum constricted, narrow, elon-
gate, exceeding middle of article 2 on antenna 1.
Eyes large, clear of pigment or deeply stained, om-
matidia ordinary. Article 1 on peduncle of antenna
1 about as long as wide, about 1.8 times as wide as
article 2, ventral margin with about 9 setules,
weakly produced dorsal apex with 3 setules; article
NUMBER 245 77
XI
FIGURE 27.?Booranus tikeri, new species, holotype, female "f," 6.92 mm; (g = male "gg," 7.4
mm; v ? male "v," size unknown).
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
k\Y
FIGURE 28.?Booranus tikeri, new species, holotype, female "f," 6.92 mm (g ? male "gg," 7.4
mm; k ? male "k," 6.60 mm; m ? male "m," 7.0 mm).
NUMBER 245 79
2 about as long as article 1, with ventral cycle of 12
setae; primary flagellum with 13 articles, about 1.1
times as long as peduncle, bearing medium aesthe-
tascs; accessory flagellum with 13 articles. Article 3
of antenna 2 with 4 setae-spines and one setule;
spine formula of article 4 = 4-3-9-9-8, dorsal mar-
gin with 4 notches each bearing 3-5 setae, ventral
margin with 6-7 groups of 1-6 long to medium
setae, 2-3 distal long spines, and clump of 11 +
setae; article 5 about 0.57 times as long as article 4,
facial spine formula = 0, dorsal margin naked or
bearing 2 setae, ventral margin with 7 unpaired
setae, clump of 5 ventrodistal long to medium spines,
no subdistal facial spines; flagellum about 1.1 times
as long as articles 4-5 of peduncle combined, with 16
articles. Epistome with sharp anterior protrusion;
upper lip with weak ventral sinus. Mandibles with
weak palpar hump; right incisor with 4 teeth; left
incisor with 4 teeth in 2 branches; right lacinia
mobilis bifid, distal branch much shorter than proxi-
mal, subbiful, proximal branch simple, pointed; left
lacinia mobilis with 4 teeth; right rakers 11 plus one
rudimentary; left rakers 12 plus one rudimentary;
molars elongate, conical, each molar with 4 long to
short spines, none disjunct; palp article 1 short, arti-
cle 2 with one long, one short inner apical setae and
6 other long and short inner and facial setae; article
3 about 1.1 times as long as article 2, oblique apex
with 12 spine-setae, basodorsal formula = 2 (one
short). Inner plate of maxilla 1 large, bearing one
long apicofacial pluseta, 3-4 shorter similar apical
setae; palp article 2 with 7 apical marginal spines
and 10 submarginal setae. Inner plate of maxilla 2
shorter and narrower than outer, outer without
apicolateral seta, inner with 9 medial setae. Inner
plate of maxilliped with 2 large thick apical spines,
5 apicofacial setae, 5 medial setae; outer plate with
16 medial and apical spines, one apicolateral seta;
palp articles 1-2 lacking lateral setae, medial mar-
gin of article 2 moderately setose, article 3 with 36
facial setae, 8 lateral setae in 3 sets, article 4 with 2
accessory setules, nail absent, Coxa 1 strongly ex-
panded distally, anterior margin almost straight;
main ventral setae of coxae 1-4 = 8-9-(9-8)-15,
posteriormost seta of coxae 1-3 shortest; coxa 2
beveled anteroventrally; anterior and posterior mar-
gins of coxa 4 divergent, posterior margin oblique,
almost straight, posterodorsal corner sharp-rounded,
posterodorsal margin short, ordinary, width-length
ratio of coxa 4 = 34:39; gills present on coxae 2-7.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = 6-5-5-6, long anteriors = 14
( + 4 facial)-6-8-l, short anteriors = 3-5-7-5, no
others. Gnathopods thin, hands thin, parachelate,
mitellid, gnathopod 2 slightly smaller than gnatho-
pod 1; width ratios of articles 5-6 of gnathopods
1-2 = 29:24 and 28:24, length ratios = 67:48 and
65:48; palmar humps very small, palms transverse;
article 5 of gnathopods 1-2 elongate, ovate, poste-
rior margins rounded-flat. Pereopods 1-2 similar;
article 4 of pereopod 2 much stouter than article 4
of pereopod 1; article 5 with apicomedial comb on
pereopods 1-2; facial setae formula on article 4 = 5
and 4, on article 5 = 5 and 6; main spine of article
5 extending to M. 75-80 on article 6, spine formula
of article 6 = 6 + 10 and 7 + 1 0 plus huge mid-
distal spine, some spines especially long; acclivity on
inner margin of dactyls of pereopods 1-2 sharp, pro-
duced as tooth, emergent setule short, marginal
pluseta ordinary. Coxae 5-7 posteroventral setae
formula = 19-19-16, some facial. Articles 4-5 of
pereopods 3-4 broad, facial spine rows dense; facial
ridge formula on article 2 of pereopods 3-5 = 0-1-1,
width ratios of articles 2, 4, 5, 6 of pereopod 3 =
50:46:41:20, of pereopod 4 = 70:55:33:14, of pereo-
pod 5 = 60:9:7:4, length ratios of pereopod 3 = 73:
40:37:34, of pereopod 4 = 75:70:47:54, of pereopod
5 = 88:20:17:20; article 2 of pereopod 5 almost
reaching apex of article 5, ventrally setose, anterior
margin weakly bulging, pereopod 5 small and
mostly hidden by pereopod 4, lateral apex of article
5 not combed; medial apex of article 6 finely combed.
Posteroventral corner of epimeron 1 rounded-
quadrate, posterior margin straight, weakly setose
(14), anteroventral margin and face with 6-8 short
setae, posteroventral margin with 1-2 short setae, no
facial setae, posterior margin with 13 setae; postero-
ventral corner of epimeron 2 rounded, posterior
margin straight, weakly serrate, setose (12-13), facial
setae = 7, none set vertically; posteroventral corner
of epimeron 3 rounded, weakly protuberant, poste-
rior margin almost straight, weakly serrate, setose
(10-11), ventral margin with 6 setae mainly in pos-
terior half, face with oblique row of 9-11 spines in
middle; epimera 1-2 with setule on posterodorsal
margin set in weak notch. Urosomite 1 with mid-
lateral group of 9-10 setae, one ventral seta on each
side, articulation line complete; urosomite 3 protu-
berant dorsally, with 2 middorsal spines on each
side. Rami of uropods 1-2 with articulate enlarged
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
sharp apical nails, outer and inner rami of uropods
1-2 with accessory nails; spike on uropod 1 reaching
to M. 55-65 on outer ramus, outer ramus of uropod
1 with 11 dorsal spines including apicals, inner with
5 and facial row of 4, not including one apical nail;
outer ramus of uropod 2 with 7 dorsal spines, in-
cluding apicals, inner with 3 dorsomedial spines in-
cluding accessory nails; peduncle of uropod 1 with
4 apicolateral spines, no basofacial setae, medially
with 2 apical spines; peduncle of uropod 2 with 6
dorsal spines, medially with 2 apical spines, apico-
lateral corners on peduncles of uropods 1-2 with
weak comb, outer rami also weakly combed. Pedun-
cle of uropod 3 with 11 + ventral spines, dorsally
with 4-5 lateral spines, one medial spine and setule;
rami masculine, inner extending to M. 100 on
article 1 of outer ramus, apex with 3 setae, medial
and lateral margins setose, article 2 of outer ramus
ordinary, 0.17, bearing 2 long setae and one me-
dium spine, apicomedial margin of article 1 setose,
lateral margin with 5 acclivities, spine formula =
1-2-2-2-2-2, setal formula = 0. Telson ordinary,
length-width ratio = 1:1, not fully cleft, each apex
wide, rounded, with or without middle acclivity,
lateral acclivity narrow, weak, bearing ordinary
lateral spine, spine next medial longer, with lateral
and medial spines separated by setule, no lateral
spine, midlateral setules alike. Cuticle with bulbar
setules closely packed with pipes and spicules,
setules not surrounded by cordate plaques, surface
bearing fine striations in form of linear fingerprint
pattern, emergent setules branched.
DESCRIPTION OF MALE.?Eyes enlarged. Article 1
of antenna 1 with 15 ventral setules, no medial
fuzz; article 2 with 15 ventral setae; primary flagel-
lum with 14 articles, one calceolus each on articles
1-5 or 1-6 or 2-6, aesthetascs on primary flagellum
as in females; accessory flagellum with 11 articles.
Article 3 of antenna 2 with facial seta-spine and
setule formula of 4 + 1; on article 4 = 4-(2+7)-9-9
or 4-(2+7)-9-8 or 4-(2 + 7)-9-7, ventral margin
with 6 sets of shortened setae; article 5 with 4-5
sets of dorsal male setae and 3 calceoli, ventrally
with 8+ setae and 2-4 apicoventral setules, flagellar
formula = 28, 2, 4, 6, . . . 26, or 36, 2, 4, 6 . . . 26,
often apical calceoli rudimentary from article 22
onwards. Palp article 3 of mandible with inner seta
near apex. Coxa 4 similar to that of female but
smaller in relation to coxa 1. Hands of gnathopods
slightly thinner than in female. Article 4 of pereo-
pods 1-2 thinner than in female. Article 2 of
pereopods 3-4 much thinner than in female. Other
highly slender articles = article 4 of pereopods 3-4;
slightly thinner = article 5 of pereopods 3-4 and
article 2 of pereopod 5; coxae 5-7 smaller relative
to their legs than in female. Epimera 1-2 broad-
ened, posterior margin of epimeron 3 not short-
ened; setal and spine formulas = epimeron 1
anteroventral = 4-6, posterior = 9, ventral = 7-9
widely spread, epimeron 2 facial = 2-5, posterior =
9-12, epimeron 3 posterior = 8-9, facial = 9-15,
ventral = 5-6. Spine formulas of uropods, uropod
1 peduncle apicolateral = 3?4, uropod 2 dorsal =
6; following notations not including apical nail
and accessory nail, dorsal spines on outer ramus of
uropod 1 = 8-9, of uropod 2 = 7-9, inner ramus
of uropod 1 * 5-6 + 3-4, of uropod 2 = 3, ventral
and lateral spines on peduncle of uropod 3 = 17,
spine formula on article 1 of outer ramus (maxi-
mum observed), spines = O-O-O-O-2-2-2-2-2-2-2,
setae = 1-1-1-2-1-1-1-1-1-1-1, spines short. Tel-
son scarcely elongate, each lobe with widely scat-
tered patch of dorsal denticles, distal spines short-
ened. Cuticular setules sparse.
OBSERVATIONS.?Male: Eyes becoming as large
as shown for male II. weemus; terminal males with
anterior spines on article 5 of pereopod 5 absent.
Juveniles: Juvenile "j," 3.95 mm, each molar
with 4 spines. Juvenile "p," 3.45 mm, left molar
with 3 teeth, right with 3 and incipient fourth.
ILLUSTRATIONS.?Maxilla 2 and maxilliped gener-
ally as shown for Tipimegus thalerus; all other
parts similar to those shown for B. weemus except
as specified in the description and attendant fig-
ures.
HOLOTYPE.?NMV, female "f," 6.92 mm.
TYPE-LOCALITY.?CPBS B6/4, 12 Oct 1964,
Western Port, Victoria, Australia, depth and sedi-
ment unknown.
VOUCHER MATERIAL.?CPBS 26N/1171: male
"k," 6.60 mm (illus.); male "m," 7.0 mm (illus.).
CPBS 25N/1171: male "gg," 7.4 mm {illus.). CPBS
B4/4: male "v," (illus.). CPBS 34N/4: juvenile "j,"
3.95 mm; juvenile "p," 3.45 mm.
RELATIONSHIP.?This species differs from the
type-species of Booranus, B. weemus, in the longer
spike on uropod 1, the broader apices of the
telsonic lobes, the fewer accessory setules on the
dactyl of the maxilliped, the presence of facial setae
on coxae 6-7, the shorter spines on the outer ramus
NUMBER 245 81
of uropod 3, the presence of 2 (not 1) spines on
each side of urosomite 3 at the base of the telson,
the absence of a lateral comb on article 5 of pereo-
pod 5, the shorter article 6 of pereopod 5, the
absence of plaques around the cuticular setules,
and in the presence of an extra set of spines on
article 4 of antenna 2.
MATERIAL.?CPBS, 20 samples from 12 stations
(155); WPBES, 5 samples from 3 stations (10);
PPBES, 2 samples from 2 stations (2).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 3-18 m, sand, sand and shell, sand
and mud, seagrass.
Boorantis wangoonis, new species
huKt 29
DESCRIPTION OF FEMALE.?Head about 23 per-
cent of total body length, greatest width about 63
percent of length, rostrum constricted, narrow
elongate, exceeding middle of article 2 on antenna
1. Eyes medium, largely occluded with pigment.
Article 1 on peduncle of antenna 1 about 1.2 times
as long as wide, about 1.8 times as wide as article 2,
ventral margin with about 7 setules, weakly pro-
duced dorsal apex with 3 setules; article 2 about 0.85
times as long as article 1, with ventral crescent of
9 setae; primary flagellum with 8 articles, about
0.85 times as long as peduncle, bearing long
aesthetascs; accessory flagellum with 8 articles
Spine formula of article 4 on antenna 2 = 2-2-6-
6-6, dorsal margin with 2 notches each bearing
one long and 1-2 short setae, ventral margin with
5 groups of 1-3 long to medium setae, dense ventro-
distal brush of 11+setae; article 5 about 0.56 times
as long as article 4, facial spine formula = 0, dorsal
margin bearing 2 setae, ventral margin with 5 sets
of one long seta each, 4 ventrodistal short to me-
dium spine-setae; flagellum about 1.1 times as long
as articles 4-5 of peduncle combined, with 11 arti-
cles. Epistome with sharp anterior protrusion;
upper lip with weak ventral sinus. Mandibles with
weak palpar hump; right incisor with 4 teeth; left
incisor with 4 teeth in 2 branches; right lacinia
mobilis bifid, distal branch little shorter than proxi-
wR3
FIGURE 29.?Booranus wangoorus, new species, holotype, male "y," 4.30 mm (u> = female "w,"
3.81 mm).
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
mal, subbifid, proximal branch simple, pointed, with
marginal denticles or facial humps; left lacinia
mobilis with 4 teeth, middle teeth shortened; right
rakers 8; left rakers 10; molars elongate, conical, each
molar with 3 (rarely 4) long to short spines, one dis-
junct, middle one completely fused to molar; palp
article 1 short, article 2 with one long inner apical
seta and one other short inner seta and 2 tiny facial
setae, article 3 about as long as article 2, oblique
apex with 9 spine-setae, basodorsal formula = one
long. Inner plate of maxilla 1 large, thin, bearing
one long apicofacial pluseta, 3 shorter apical setae;
palp article 2 with 7 apical marginal spines and 8
submarginal setae (4 of these facial). Inner plate of
maxilla 2 shorter and narrower than outer, outer
with one apicolateral seta, inner with 3 medial
setae. Inner plate of maxilliped with 2 large thick
apical spines, 5 apicofacial setae, 3 medial setae;
outer plate with 11 medial and apical spines, one
apicolateral seta; palp articles 1-2 without apico-
lateral setae, medial margin of article 2 strongly
setose, article 3 with 14 facial setae, 4 lateral setae
in 4 groups, article 4 with 2 accessory setules, one
on face, nail absent. Coxa 1 strongly expanded
distally, anterior margin almost straight; main
ventral setae of coxae 1-4 = 6-7-6-11, posterior-
most seta of coxae 1-3 shortest; anterior and pos-
terior margins of coxa 4 divergent, posterior margin
oblique, almost straight, posterodorsal corner sharp,
posterodorsal margin of medium length, concave,
width-length ratio of coxa 4 = 7:8; gills present
on coxae 2-7. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 4-3-3-4,
short posteriors = 0-0-0-1, long anteriors = 13(+4
facial)-8-6-0, short anteriors = 1-1-3-2, no others.
Gnathopods thin, hands thin, parachelate, mitellid,
gnathopod 2 slightly smaller than gnathopod 1;
width ratios of articles 5-6 on gnathopods 1-2 =
26:22 and 25:21, length ratios = 64:42 and 61:42;
palmar humps very small, palms transverse; article
5 of gnathopods 1-2 elongate, ovate, posterior mar-
gin rounded-flat. Pereopods 1-2 similar, article 4
of pereopod 2 stouter than article 4 of pereopod 1;
article 5 with apicomedial comb on pereopods 1-2;
facial setae formula on article 4 = 3 and 2, on
article 5 = 3 and 4, main spine of article 5 extend-
ing to M. 80 on article 6, article 5 with one extra
long distoposterior spine and numerous setae, prox-
imal margin naked, spine formula of article 6 =
4+6 and 5+6 plus huge middistal spine, some
spines especially long; acclivity on inner margin of
dactyls of pereopods 1-2 sharp, produced as tooth,
emergent setule short, marginal pluseta ordinary.
Coxae 5-7 posteroventral seta formula = 11-12-10
(only 1-2 facial on coxa 7). Articles 4-5 of pereo-
pods 3-4 broad, facial spine rows dense; facial
ridge formula on article 2 of pereopods 3-5 =
0-1-1, width ratios of articles 2, 4, 5, 6 of pereopod
3 = 46:44:37:19, of pereopod 4 = 65:52:35 14, of
pereojxxl 5 = 70:11:9:6, length ratios of pereopod
3 = 75:3940:33, of pereopod 1 = 80:66:5055, of
pereopod 5 = 88:24:2325; article 2 of pereopod 5
reaching middle of article 5, ventrally setose, ante-
rior margin scarcely bulging, pereopod 5 small and
mostly hidden by pereopod 4, lateral apex of article
5 and medial apex of article 6 finely combed.
Posteroventral corner of epimeron I rounded-
quadrate, |x>sterior margin straight, strongly setose
(11-14), anterovcntral margin and face with 5 short
setae, postcroventral margin with 2 short setae, face
with 2-3 setae near posterior margin; posteroven-
tral corner of epimeron 2 rounded, posterior mar-
gin straight, weakly serrate, setose (6-9), facial
setae = 3, none set vertically; posteroventral corner
of epimeron 3 rounded, weakly protuberant, poste-
rior margin almost straight, serrate, setose (6), ven-
tral margin with 5 setae mainly in posterior half,
face with oblique row of 7 spines in middle,
epimera 1-2 with setule on posterodorsal margin
set in weak notch. Urosomite 1 with midlateral
group of 6 setae, one ventral seta on each side,
articulation line complete; urosomite 3 protuberant
dorsally, with middorsal spine on each side. Rami
of uropods 1-2 with articulate enlarged sharp
apical nails, outer and inner rami of uropods 1-2
with accessory nails; spike on uropod 1 reaching to
M. 50 on outer ramus, outer ramus of uropod 1
with 8 dorsal spines including nails, inner with 5
including nails and facial row of one spine; outer
ramus of uropod 2 with 6 dorsal spines including
nails, inner with 3 dorsomedial spines including
nails; outer rami of uropods 1-2 very weakly
combed; peduncle of uropod 1 with 2 apicolateral
spines, no basofacial setae, medially with 2 mar-
ginal spines, apicalmost ordinary; peduncle of
uropod 2 with 2-3 dorsal spines, medially with pair
of apical spines; apicolateral corners on peduncles
of uropods 1-2 with comb. Peduncle of uropod 3
with 9 ventral spines, well segregated from one
dorsal and lateral spine, one medial spine; rami
NUMBER 245 83
submasculine, inner extending to M. 100 on article
1 of outer ramus, apex with 3 setae, medial and
lateral margins setose; article 2 of outer ramus
ordinary, 0.26, bearing 2 long setae and one spine,
apicomedial margin of article 1 setose, lateral mar-
gin with 3-4 acclivities, spine formula = 2-2-2-2
or 1-2-2-2-2; setal formula = 0. Telson ordinary,
length-width ratio = 1:1, not fully cleft, each apex
narrow, rounded, with middle acclivity, lateral
acclivity narrow, weak, tearing ordinary lateral
spine, spine next medial subequal, lateral and
medial spines separated by setule, midlateral setules
diverse. Cuticle with bulbar setules mixed with
pipes and spicules, setules not surrounded by
cordate plaques, surface tearing fine striations in
form of linear fingerprint pattern, emergent setules
branched.
OBSERVATIONS (female).?Description based on fe-
male "w"; male "y" fully described below owing to
remote possiblity these two specimens not conspeci-
fic. Article S on peduncle of antenna 2 with groups of
3 setae and one setule. Spines on outer ramus of
uropod 3 almost as long as in Booranus weemus;
coxae 1-3 slightly larger relative to coxa 4 than in
B. weemus; eyes slightly smaller than B. weemus;
inner plate of maxilla 1 thinner than in B. weemus;
article 4 of pereopods 1-2 about 15 percent thinner
and article 6 about 10 percent shorter than in B.
weemus.
DESCRIPTION OF MALE.?Head about 21 percent
of total body length, greatest width about 66 per-
cent of length, rostrum constricted, narrow, elon-
gate, exceeding middle of article 2 on antenna 1.
Eyes large, largely occluded with pigment. Article 1
on peduncle of antenna 1 about 1.1 times as long as
wide, about twice as wide as article 2, ventral mar-
gin with about 13 setules, weakly produced apex
with 3 setules; article 2 about as long as article 1,
with ventral crescent of 9 setae; primary flagellum
with 8 articles, about 0.85 times as long as peduncle,
bearing long aesthetascs; accessory flagellum with 8
articles. Spine formula on article 4 of antenna 2 =
2-2-6-7-5, dorsomedial margins of articles 3-4
fuzzy, ventral margin with 6 groups of 1-3 long to
medium setae, ventrodistal brush of 7-12 either all
long or all short setae; article 5 about 0.77 times as
long as article 4, facial spine formula = 2 dorso-
distal, dorsal margin tearing 4 groups of male setae
and 3 calceoli, ventral margin with 5 sets of one
short seta each, 3 ventrodistal setules set facially;
flagellar formula = 27, 2, 3, 5, 7 . . . 19. Epistome
with sharp anterior protrusion; upper lip with weak
ventral sinus. Mandibles with weak palpar hump;
right incisor with 4 teeth; left incisor with 4 teeth
in 2 branches; right lacinia mobilis bifid, distal
branch shorter than proximal, subbifid, denticulate,
with 2 facial humps, proximal branch simple blunt,
with marginal denticles or facial humps; left lacinia
mobilis with 4 teeth, middle teeth shortened; right
rakers 7 plus one rudimentary; left rakers 9 plus one
rudimentary; molars elongate, conical, each molar
with 3 long to short spines, one disjunct, mildle one
completely fused to molar; palp article 1 short,
article 2 with one long inner apical seta and 2 other
short inner setae and one medium facial seta, article
3 about as long as article 2, oblique apex with 10
spine-setae, basofadal formula = 2 (one long, one
very short). Inner plate of maxilla 1 large, thin,
bearing one long apicofacial pluseta, 3 shorter
apical setae; palp article 2 with 8-9 apical marginal
spines and 8-10 submarginal setae (4-6 facial). In-
ner plate of maxilla 2 slightly shorter and narrower
than outer, outer with one apicolateral seta, inner
with 5 medial setae. Inner plate of maxilliped with
2 large thick apical spines, 5 apicofacial setae, 3
medial setae; outer plate with 13 medial and apical
spines, one apicolateral seta; palp articles 1-2 with-
out apicolateral seta, medial margin of article 2
strongly setose, article 3 with 17 facial setae, 5
lateral setae in 4 groups, article 4 with 2 accessory
setules, nail absent. Coxa 1 strongly expanded
apically, anterior margin almost straight; main ven-
tral setae of coxae 1-4 = 6-8-7-13, posteriormost
seta of coxae 1-3 shortest; anterior and posterior
margins of coxa 4 divergent, posterior margin
oblique, almost straight, posterodorsal corner sharp,
posterodorsal margin concave, undulant, width-
length ratio of coxa 4 = 33:38; gills present on
coxae 2-7. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 3-3-4-4, short
posteriors = 0, long anteriors = 17(+3 facial)-9-
8-1, short anteriors = 1-2-2-2, no others. Gnatho-
pods thin, hands thin, parachelate, mitellid, gnatho-
pod 2 slightly smaller than gnathopod 1; width
ratios of articles 5-6 on gnathopods 1-2 = 25:21
and 27:20, length ratios = 64:42 and 62:41; palmar
humps small, palms transverse; article 5 of gnatho-
pods 1-2 elongate, ovate, posterior margin rounded-
flat. Pereopods 1-2 similar, article 4 even thinner
than in female, but stouter on pereopod 2 than on
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pereopod 1; article 5 with apicomedial comb, facial
setae formula on article 4 = 4 and 2, on article
5 = 4 and 4, main spine of article 5 extending to
M. 77 on article 6; article 5 with one extra long
distoposterior spine and numerous setae, proximal
margin naked; spine formula of article 6 = 5 + 6
and 5 + 6 plus huge middistal spine, some spines
especially long; acclivity on inner margin of dactyls
of pereopods 1-2 sharp, produced as tooth, emer-
gent setule short, marginal pluseta ordinary. Coxae
5-7 posteroventral seta formula = 13-13-11 (only
one facial on coxa 7). Articles 4-5 of pereopods 3-4
broad, facial spine rows dense, facial ridge formula
on article 2 of pereopods 3-5 = 0-1-1, width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 43:40:38:18, of
pereopod 4 = 58:47=28:11, of pereopod 5 = 65:
10:9:4, length ratios of pereopod 3 = 68:37:38:30,
of pereopod 4 = 75:65:44:50, of pereopod 5 =
90:21:22:23; article 2 of pereopod 5 exceeding
middle of article 5, ventrally setose, anterior margin
scarcely bulging, pereopod 5 small and mostly hid-
den by pereopod 4, lateral apex of article 5 and me-
dial apex of article 6 finely combed. Epimera 1-3
broadened; posteroventral corner of epimeron 1
rounded-quadrate, posterior margin almost straight,
strongly setose (14), anteroventral margin with 6-7
short setae, posteroventral margin with 5 medium
setae, one fully facial short seta; posteroventral cor-
ner of epimeron 2 rounded, weakly protuberant,
with seta notch, posterior margin straight, setose
(10), facial setae = 3, none set vertically; postero-
ventral corner of epimeron 3 rounded, weakly pro-
tuberant, posterior margin straight to weakly convex,
not shortened, weakly serrate, setose (7), ventral mar-
gin with 5 setae mainly in posterior half, face with
oblique row of 7 spines in middle; epimera 1-2 with
setule on posterodorsal margin set in weak notch.
Urosomite 1 with midlateral group of 7 setae, one
ventral seta on each side, articulation line incom-
plete, short; urosomite 3 protuberant dorsally, with
middorsal spine on each side. Rami of uropods 1-2
with articulate enlarged sharp apical nails, spike on
uropod 1 reaching to M. 50 on outer ramus, outer
and inner rami of uropods 1-2 with accessory nails,
outer ramus of uropod 1 with 10 dorsal spines
including nails, inner with 6 including nails plus
one in facial row, outer ramus of uropod 2 with 9
dorsal spines including nails, inner with 6 dorso-
medial spines including nails, outer rami of uro-
pods 1-2 very weakly combed; peduncle of uropod
1 with 2 apicolateral spines, no basofacial setae,
medially with 3 marginal spines, apicalmost ordi-
nary; peduncle of uropod 2 with 5 dorsal spines,
medially with 2 spines, apicolateral corners on
peduncles of uropods 1-2 with comb. Peduncle of
uropod 3 with 13 ventral spines, dorsally with one
lateral spine, one medial spine and setule; rami
masculine, inner extending to M. 120+ on article 1
of outer ramus, apex with 3 setae, medial and lateral
margins setose, article 2 of outer ramus short, 0.21,
bearing 2 long setae, one spine, occasionally one
setule, medial margin of article 1 setose, lateral mar-
gin with 6 acclivities, spine formula = 0-0-2-2-2-2-
2, spines medium, setal formula = 1-2-1-1-1-1-1.
Telson long, length-width ratio = 15:11, not fully
cleft, each apex narrow, rounded, with middle ac-
clivity, lateral acclivity narrow, weak, bearing ordi-
nary lateral spine, spine next medial longer, with
lateral and medial spines separated by setule, mid-
lateral setules equal to each other. Cuticle with
bulbar setules mixed with pipes and spicules, setules
not surrounded by cordate plaques, surface bearing
fine striations in form of linear fingerprint pattern,
emergent setules branched.
OBSERVATIONS (male).?Description based on male
"y." Article 5 of gnathopod 1 slightly thinner than
in female, article 6 slightly shorter, palmar hump
slightly larger; article 3 on peduncle of antenna 2
with groups of 3 setae and one setule; antenna 2
more strongly ensiform than in female or in Boora-
nus weemus; article 5 of pereopod 5 with 4 ante-
rior spines and setae; pereopods 3-4 smaller relative
to pereopod 5 than in female of B. weemus; coxa 4
about as shown for female of B. weemus; coxa 1
not differentially enlarged relative to ratios of coxae
1 and 4 in female of B. weemus.
VARIATIONS.?Females "k," "m," and "n" with 5
setae on inner plate of maxilla 1; female "n" with
poorly developed fourth tooth on molar, speci-
men otherwise not senile; female "k" with only 7
rakers on each mandible, bearing incipient fourth
molarial tooth; female "m," though small, bearing
full 12 apical setae on article 3 of mandibular palp,
right rakers 10 plus one rudimentary, left rakers 11
plus one rudimentary; largest ovigerous female in
collections, 5.85 mm, and parts mentioned for fe-
male "m" identical; one subadult male from CPBS
01N with incipient fourth tooth on molar; juve-
NUMBER 245 85
nile, 3.75 mm, with 5 teeth on left lacinia mobilis,
only 3 molarial teeth, 9 apical spines on palp arti-
cle 3 of mandible and 8 raker spines (left side only
observed).
ILLUSTRATIONS.?Parts conforming to those of
Booranus weemus except as noted in description
and observations.
HOLOTYPE.?NMV, male "y," 4.30 mm.
TYPE-LOCALITY.?PPBES 904/5, 7 Jun 1971, Port
Phillip Bay, Victoria, Australia, 9 m, silty sand.
VOUCHER MATERIAL.?Type-locality, female "w,"
3.81 mm (illus.); CPBS 10E/2, female "k," 5.05 mm;
CPBS 25S/3, female "m," 4.42 mm; CPBS 01N/3,
female "n," 5.00 mm.
RELATIONSHIP.?This species is extremely difficult
to separate from the other two species of Booranus.
It is a dwarf species recognized by its small size at
maturity, about 4.0 mm in body length. Specimens
of the other species in that size range are juveniles
with greatly reduced setation on epimera compared
with 4.0 mm adults of B. wangoorus. It is easily
differentiated from B. weemus in the longer spike
on uropod 1 that reaches M. 50 on the outer
ramus. The terminal males are easily recognized
because they bear 2 facial spines on article 5 of
antenna 2 in contrast to one spine in the other
species. The mandibular molar almost never bears
more than 3 tooth-cusps in. specimens of 4.0 mm
body length whereas juveniles of the other species
bear 4 tooth-cusps, although specimens of those
other species 3.5 mm long have only 3 teeth or 3
plus one rudimentary. In some respects the species
is closer to B. weemus than to B. tikeri: article 5
of pereopod 5 is combed apicolaterally and uroso-
mite 3 bears only one dorsal spine on each side in
contrast to the two spines on each side in B. tikeri.
The broadened telson of B. tikeri is usually distinc-
tive though not easy to estimate in undissected
view. Article 2 on the outer ramus of uropod 3 is
somewhat more elongate in B. wangoorus than in
the other two species, and unlike B. tikeri, coxa 3
is not beveled anteroventrally. The apical spines
on each lobe of the telson are somewhat more
widely spread in B. wangoorus than in the other
two species. Unlike B. tikeri, coxa 7 has one facial
seta in B. wangoorus.
MATERIAL.?PPBES, 21 samples from 8 stations
(77); CPBS, 21 samples from 12 stations (31);
WPBES, 4 samples, from 3 stations (24); AM, 2
samples (5).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 0-14 m, sand, silty sand, silty clay,
sand and shell, seagrass. South Australia: Kangaroo
Island, neritic. Bass Strait: 70 miles south of Lakes
Entrance, 95 m.
Trichophoxus K. H. Barnard
Trichophoxus K. H. Barnard, 1930:336.?Barnard and Drum-
mond, 1976:534.
DIAGNOSIS OF MALE.?Eyes present. Flagella of
antennae 1-2 unreduced in male. Article 2 of an-
tenna 1 elongate, ventral setae widely spread. Arti-
cle 1 of antenna 2 weakly ensiform, article 3 with
numerous setae and setules, facial spines on article
4 in 2 or more rows plus special apical spines, arti-
cle 5 short. Right mandibular incisor with 4 teeth;
molar not triturative, large, elongate-conical, then
subtruncate and bearing 4 special large spines, not
bearing fuzz; palpar hump small. Palp of maxilla
1 biarticulate, inner plate with 4 setae. Setation of
maxilla 2 ordinary. Inner plate of maxillipeds ordi-
nary; apex of palp article 3 not protuberant, dactyl
elongate, apical nail short, mostly immersed. Gills
present only on coxae 2-6. Gnathopods small, simi-
lar, article 5 of gnathopods 1-2 free, elongate, with-
out eusirid attachment; palms transverse, almost
chelate, hands of gnathopods 1-2 elongate, heavily
setose anteriorly, trichophoxid in shape. Article 5
of pereopods 1-2 lacking posteroproximal setae.
Article 2 of pereopod 3 of broad form, articles 4?5
of pereopods 3-4 broad, article 2 of pereopods 3-4
weakly setose posteriorly, pereopod 5 especially
small, article 2 strongly setose ventrally, article 3
greatly enlarged, dactyl normal. Epimera 1-2 bear-
ing numerous long posterior setae, with midfacial
setae above ventral facial ridge, epimeron 3 ordinary;
urosomite 1 with large lateral facial spines, bearing
one or more midventral crescents or bundles of setae,
urosomite 3 without dorsal hook or special process.
Peduncle of uropod 1 normally elongate, with apico-
ventral small spike, without special enlarged apico-
lateral-medial spine, peduncular apices of uropods
1-2 combed, inner ramus of uropod 1 with one row
of marginal spines, only one ramus continuously spi-
nose to apex, inner ramus of uropod 2 ordinary
but naked. Uropod 3 ordinary, article 2 of outer
ramus carrying 2-3 long apical setae. Telson with
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
2 apical spines on each lobe plus setules, with spe-
cial lateral spines.
DESCRIPTION OF MALE.?Rostrum constricted,
small. Fuzz on article 1 of antenna 1 in male absent;
calceoli on male primary flagellum of antenna 1
present. Calceoli on article 5 of male antenna 2
present, flagellum in male with calceoli. Prebuccal
parts extended forward, especially massive, strongly
distinct, upper lip dominant. Right lacinia mobilis
bifid, thin; article 1 of mandibular palp short, palp
thin, apex of article 3 oblique, article 2 with outer
setae. Lower lip lacking cones. Outer plate of maxilla
1 with 11 spines, one spine especially thickened.
Inner plates of maxilliped thick, ordinarily setose.
Coxae 2-4 with special anterodorsal humps. All
posterior spines on article 6 of pereopods 1-2 thick
and stiff, midapical spine or seta present, especially
enlarged. Article 2 of pereopod 5 with facial setae.
Peduncle of uropod 1 with dorsolateral spines
widely spread, of uropods 1-2 with medial spines
widely spread. Peduncle of uropod 3 lacking extra
subapical setae or spines. Telson with midlateral
setules highly apical, reduced to one member on
each side.
TYPE-SPECIES.?Trichophoxus capillatus K. H.
Barnard, 1930 (monotypy).
COMPOSITION.?Unique.
RELATIONSHIP.?Trichophoxus is distinguished
from Tipimegus in numerous attributes, among
them: (1) lateral spination on urosome; (2) lateral
spination on telson; (3) far more numerous setae
on article 4 of antenna 2; {4) presence of 4 setae
(not 3) on inner plate of maxilla 1; (5) immensity
of apical midspine on article 6 of pereopods 1-2;
(6) elongation and bareness of inner ramus on uro-
pod 2; (7) normally elongate peduncle of uropod
1; (8) proximal spination on peduncle of uropod 1;
(9) presence of only one row of spines on inner
ramus of uropod 1; (10) presence of only one ven-
tral crescent of setae on urosomite 1; (11) strong
distal shift of and reduction in number of dorsal
setules on telson; (12) absence of any combs on
pereopods 3-5; (13) presence of facial humps on
right lacinia mobilis; (14) distad shift in primary
tooth on dactyls of pereopods 1-2; (15) presence
of more than one medial spine on peduncle of
uropod 2.
Trichophoxus is more primitive than Tipimegus
in such characters as peduncles of uropods 1-2 and
lack of pereopodal combs but is far more special-
ized than the Australian genus in telsonic and uro-
somal ornamentation.
Trichophoxus capillatus K. H. Barnard
Trichophoxus capillatus K. H. Barnard. 1930:336-357, fig.
13? Barnard and Drummond. 1976:536-543, hgi. 1-3, 4
[part].
DISTRIBUTION.?New Zealand, North Cape, 3 m,
neritic.
Waitangi Fincham
Waitangi Fincham, 1977:286.
DIAGNOSIS (based on literature prior to Fincham,
1977).?Eyes present. Flagella of antennae 1-2 un-
reduced in female. Article 2 of antenna 1 ordinary,
ventral setae widely spread. Article 1 of antenna 2
weakly ensiform; article 3 with numerous setae and
setules; facial spines on article 4 in 2 or more rows
[?plus special apical spines]; article 5 thin and
short. [Right mandibular incisor with ?4 teeth];
molar not triturative, large, elongate-conical, then
subtruncate and bearing 4 special large spines, not
bearing fuzz; palpar hump small. Palp of maxilla
1 biarticulate; [inner plate with ? 2 3 4 5 + setae.
Setation of maxilla 2 ordinary, ?weak. Inner plate
of maxilliped Pordinary, Pmostly fused together;
Papex of palp article 3 not or weakly strongly pro-
tuberant, Pdactyl elongate, short, stubby, Papical
nail distinct, mostly immersed, short, medium,
elongate, obsolescent, absent]. Gnathopods similar;
article 5 of gnathopods 1-2 free, elongate, without
eusirid attachment; palms transverse to chelate;
hands of gnathopods 1-2 ordinary, elongate, heav-
ily setose anteriorly, trichophoxid in shape. Article
5 of pereopods 1-2 lacking posteroproximal setae.
Article 2 of pereopod 3 of broad form; articles 4-5
of pereopods 3-4 broad; article 2 of pereopods 3-4
setose posteriorly; pereopod 5 especially small, arti-
cle 2 strongly setose ventrally, article 3 greatly en-
larged, dactyl normal. Epimera 1-2 bearing long
posterior setae, with midfacial setae above ventral
facial ridge; epimeron 3 ordinary, with large tooth.
Urosomite 1 [with Plarge lateral facial spines], bear-
ing one or more midventral crescents or bundles of
setae; urosomite 3 without dorsal hook or special
process. Peduncle of uropod 1 normally elongate,
with apicoventral small spike, without special en-
NUMBER 245 87
larged apicolateral-medial spine; peduncular apices
of uropods 1-2 not combed, but peduncles and
rami with long setae, inner ramus of uropod 1 with
one row of marginal spines, all rami continuously
spinose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, article 2 of outer ramus [carry-
ing ?one, 2, 3 long vestigial apical setae]. Telson
ordinary, with 2-4 apical spines or setae on each
lobe plus setules.
DESCRIPTION.?Rostrum constricted, small. [Fuzz
on article I of antenna 1 in male unknown, cal-
ceoli on male primary flagellum of antenna 1 un-
known, calceoli on male antenna 2 unknown]. Pre-
buttal parts strongly extended forward, especially
massive, strongly distinct, both epistome and upper
lip dominant. [Right lacinia mobilis Phiful, Pflabel-
late]; article 1 of mandibular palp short, palp thin,
apex of article 3 oblique, article 2 with outer setae;
[?Lower lip bearing, lacking cones. POuter plate of
maxilla 1 with 7, 9, 11 spines, no or one spine espe-
cially thickened. Pinner plates of maxilliped espe-
cially thin, poorly armed, thick, ordinarily setose].
Coxae 2-4 with special anterodorsal humps. All
posterior spines on article 6 of pereopods 1-2 thin
and stiff, midapical spine or seta present, especially
enlarged. Article 2 of pereopod 5 with facial setae.
Peduncle of uropod 1 with dorsolateral spines con-
fined apically, medial spines confined apically; pe-
duncle of uropod 2 with medial spines widely
spread. Peduncle of uropod 3 lacking extra sub-
apical setae or spines. Telson with midlateral or
dorsal setules on each side highly apical, reduced
to one member on each side.
TYPE-SPECIES.?Paraphoxus rakiura Cooper and
Fincham, 1974.
COMPOSITION.?Unique.
RELATIONSHIP.?Waitangi resembles Tricho-
phoxus more than it does Tipimegus or Booranus
in the elongate peduncle of uropod 1, elongate
midapical spine on article 6 of pereopods 1-2,
smallness of spike on uropod 1, and presence of
facial setae on article 2 of pereopod 4, but resem-
bles Tipimegus more in the presence of spine-setae
on the inner ramus of uropod 2, the narrow spread
of apicolateral spines on the peduncle of uropod 1,
in the absence of lateral spines on the telson, and
the lack of lateral spines on urosomite I.
Waitangi differs from both Trichophoxus and
Tipimegus in the presence of long setae on the
peduncles and rami of uropods 1-2 and in the
.huge tooth of epimeron 3, a radical development
in the tipimegin group.
Waitangi rakiura (Cooper and Fincham)
Paraphoxus rakiura Cooper and Fincham, 1974:168-173,
figs. 7b, 8-10.
Waitangi rakiura.?Fincham, 1977:286.
DISTRIBUTION.?New Zealand, Stewart Island, in-
tertidal sand.
BROLGINAE
DIAGNOSIS.?Article 2 of antenna 1 shortened;
mandibular molar nontriturative, bearing 3 or
fewer tightly clumped spines; palp of maxilla 1
biarticulate; setation on maxilla 2 ordinary to re-
duced; gnathopod 2 either enlarged or not; article
2 of pereopod 3 of broad form, pereopod 5 with
ordinary article 3.
DESCRIPTION.?Article 5 of antenna 2 reduced;
epimeron 3 of rounded classification; apices of pe-
duncles on uropods 1-2 usually combed.
TYPE GENUS.?Brolgus, new genus.
COMPOSITION.?Cunmurra, new genus (placed
here provisionally, technically assignable to Joubi-
nellinae), Elpeddo, new genus, Ganba, new genus,
Kuritus, new genus, Mandibulophoxus J. L. Bar-
nard (satellite genus), Paraphoxus Sars, and Wil-
dus, new genus.
THE BROLGIN GROUP OF GENERA
Brolgus tattersalli (J. L. Barnard) and nine other
new species from Australia constitute the brolgin
group of genera, divided into Brolgus, Elpeddo,
Ganba, Kuritus, and Wildus. A sixth genus, Para-
phoxus, now restricted to one or two species, forms
the ultimate evolute confined to north boreal wa-
ters. A seventh genus, Cunmurra, forms an inter-
gradation between the brolgin.group and the Bi-
rubius mayamayi group of species. Cunmurra has
the gnathopods and epimera of a brolgin but has
one too many spines (4) on the mandibular molar
and far too elongate an article 2 of antenna 1 to
qualify as a fully fledged brolgin. Mandibulophoxus
is a satellite genus perhaps deserving a subfamily of
its own.
One main feature of the brolgin group in its
primitive state is the diversity between gnathopods
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
1 and 2 and the enlargement of gnathopod 2. This
state is fully expressed in Brolgus, where gnatho-
pod 2 is very large, the wrist is short, and the
strongly oblique palm is almost as long as or longer
than the posterior margin of the hand. The hand
of gnathopod 1 is generally thinner than in typical
Australian species of Bintbius and this becomes
even thinner in other genera of the Brolgus group.
In Kuritus, gnathopod 2 assumes the thin handed
form also.
Because of variability in gnathopods other char-
acters serve better to identify the Brolgus group.
All known genera of the Brolgus group have the
following characters in common: molars with only
3 or fewer tightly clumped large spines, molar
otherwise weak; article 2 of antenna 1 very short;
third and fourth facial rows of spines on article 4
of antenna 2 amalgamated; setae on inner plate of
maxilla 1 reduced, generally to 2 members; pereo-
pods 4?5 with only one main facial ridge on article
2, occasionally with weak second ridge; inner plate
of maxilliped with only one main distal spine, this
spine long and thin, weakly feathered or not; epi-
meron 3 of rounded classification, lacking any elon-
gate setae, protruding posteriorly and bearing at
most 3 setule notches. The thinness of spines on
article 4 of antenna 2 creates confusion between
those spines and the plumose accessory setules eas-
ily distinguished in Birubius. Proximal groups of
spines in members of the brolgin group have been
counted by subtracting the accessory setules iden-
tified by their plumosities.
The following characters also are of use in recog-
nizing members of the Brolgus group but are not
present in all genera: apical combs on peduncles
of uropods 1-2 (except in Ganba); articles 4-5 of
pereopods 3-5 very thin except in Ganba where
pereopod 3 is slightly thickened; article 5 of an-
tenna 2 shortened in females (unknown in Kuritus
because the female is yet unknown); article 6 of
pereopods 1-2 with very thin posterior spines (ex-
cept in Ganba and Elpeddo). Brolgus and Ganba
have lost one of the two apical setae on article 2
of the outer ramus on uropod 3 typical of biru-
biins. All species have an unprotuberant article 3
on the maxilliped, normal nail on the dactyl of
the maxilliped, lack accessory apical spines or con-
tinuous spination on the rami of uropods 1-2 but
generally, in other respects, conform to the birubiin
morphology.
Cunmurra, the satellite genus, possibly primitive,
has weakly enlarged gnathopod 2 but the wrist is
not fully shortened, and, in addition, Cunmurra
lacks the following features of the Brolginae: sparse
molarial spination, sparse setation on inner plate
of maxilla I, thin single apical spine on inner plate
of maxilliped, short article 2 of antenna 1 and
article 5 of antenna 2, amalgamated spine rows on
article 4 of antenna 2, thin spines on pereopods
1-2, thin distal articles on pereopods 5-5, and pres-
ence of combs on uropods 1-2. Cunmurra also bears
2 ventral setae on epimeron 3, a condition not typi-
cal of brolgins. Cunmurra thus demonstrates the
close convergence between the Birubius mayamayi
group of species and the brolgin genera but is also
a broadly transitional genus because it could be
assigned to the Joubinellinae owing to its mandib-
ular molar, elongate article 2 of antenna 1, and
enlarged gnathopod 2.
After Cunmurra, Ganba is the next weakest
member of the Brolgus group because of the ab-
sence of uropodal combs and thicker articles 4-5
of pereopod 3 but Ganba does not form a good
intergrade between Cunmurra and the other brol-
gin genera because of specializations in gnathopods
and maxillipeds. The gnathopods are thin and the
inner plates of the maxilliped have become par-
tially fused, truncate, and bear all their setae dis-
tally. This condition is, however, signaled in Wil-
dus, which has partially fused plates with mostly
terminal setae, but with the apices not fully trun-
cate. Ganba also has thickened spines on pereopods
1-2, a primitive condition in this context.
Kuritus is a poor intergrade between Cunmurra
and any other brolgin, although it maintains in
primitive condition the pair of apical setae on the
outer ramus of uropod 3, strong basofacial setae
on uropod 1, and has the more advanced spination
of pereopods 1-2. Gnathopods of Kuritus, being
thin and similar, have departed widely from the
brolgin concept.
No even-handed line of descent occurs in the
known taxa because of the mixture of characters.
One ultimate brolgin, Brolgus, has enlarged gnath-
opod 2 and only one seta on uropod 3, whereas
another, Ganba, in addition to these characters,
retains mostly short posterior spines on article 6
of pereopods 1-2 and has more specialized inner
plates on the maxillipeds. Wildus does not form a
good step towards Ganba because it combines fully
NUMBER 245 89
specialized spines on the pereopods with primitive
maxillipeds. Wildus would be considered a good
ancestral type for Brolgus except for the weak spe-
cialization in maxillipeds.
Elpeddo also retains 2 apical setae on the outer
ramus of uropod 3 but these are very short, almost
vestigial. The gnathopods are scarcely enlarged.
The male of Elpeddo, perhaps aberrantly, has de-
veloped a swollen article 1 on the primary flagel-
lum; this article bears rows of elongate aesthetascs
in the form of an enormous antennal brush. None
of the posterior spines on article 6 of pereopods
1-2 is elongate and thin.
The tropical Mandibulophoxus J. L. Barnard is
considered to be an advanced satellite of the Brol-
ginae in which the palpar hump of the mandible
is huge, the molar is reduced or absent, and on the
rami of uropods 1-2 dorsal spination is continuous
to the apex. Article 5 of antenna 2 is not shortened
relative to article 4 but the latter itself is short-
ened, suggesting that article 5 is indeed in the
shortened configuration. Article 2 of pereopod 5
is extended posteroventrally. Epimeron 3 has a few
ventral setae. Article 6 of the gnathopods is not
truly brolgin because it is expanded distally but
gnathopod 2 otherwise meets brolgin requirements
in that it is larger than gnathopod 1 and has a
cryptic wrist. Mandibulophoxus may therefore
branch away from a hypothetical brolgin precursor
rather than stand directly in the mainstream of this
group. The continuous dorsal spination on uro-
podal rami is one primitive mark suggesting this
early branching.
RELATIONSHIPS OF THE BROLGIN GROUP TO
Metaphoxus AND Phoxocephalus
The Brolgus group of genera is easily confounded
with species in Metaphoxus and Phoxocephalus
because of external resemblances. Both groups have
a poorly setose epimeron 3 with bulging posterior
margin and both have apical combs on the pe-
duncle of uropods 1-2. The palp of maxilla 1 is
uniarticulate in Metaphoxus and Phoxocephalus
but biarticulate in Brolgus. Phoxocephalus, unlike
Brolgus and Metaphoxus, has a fully triturative
molar. The molar of Brolgus bears 3 -heavy spines
and that of Metaphoxus is armed with numerous
thin setules. In addition, Brolgus has 1-2 long api-
cal setae on the outer ramus of uropod 3 which
are not present in Metaphoxus and Phoxocephalus.
If the outer ramus of uropod 1 is slightly shortened
then one is not dealing with a member of the Brol-
gus group, though only a few species of the other
genera have that ramus shortened. Several brolgins
retain well setose second maxillae in contrast to the
Metaphoxus group.
The close resemblance of the Brolgus group to
Metaphoxus and Phoxocephalus suggests a possible
direct evolutionary relationship. If one holds the
thesis that triturative molars are more primitive,
then Phoxocephalus might be ancestral to the
Brolgus group, but this would require an evolution-
ary reversal in segmentation on die palp of maxilla
1. Phoxocephalus could be, however, ancestral to
Metaphoxus, without this reversal. The Brolgus
group could be ancestral to Metaphoxus but not to
Phoxocephalus without reversal in evolution of the
molar. Metaphoxus and Phoxocephalus are so close
in many features, including the uniarticulate first
maxillary palp, that their independent evolutionary
development is not logically held. One must there-
fore conclude, provisionally, that the Brolgus group
is simply convergent, sharing the Metaphoxus and
Phoxocephalus similarities in epimera and uropods.
Key to the Genera of Brolginae
1. Article 2 of peduncle on antenna 1 elongate, mandible with 4 or more spines on molar
(not fully typical of brolgins) Cumnurra, new genus
Article 2 of peduncle on antenna 1 shortened, mandible with 3 or fewer spines on molar
2
2. Apex of outer ramus on uropod 3 with 2-3 setae S
Apex of outer ramus on uropod 3 with one seta 7
3. Rami of uropods 1-2 continuously spinose to apex Mandibulophoxus
Rami of uropods 1-2 not continuously spinose to apex 4
4. Uropod 1 lacking special apicomedial spine on peduncle Paraphoxus
Uropod 1 bearing special apicomedial spine on peduncle 5
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
5. Gnathopods identical to each other, hands thin Kuritus, new genus
Gnathopods diverse, hands weakly to broadly expanded 6
6. Apical setae on outer ramus of uropod 3 shorter than article 2, article 5 of gnathopod 2
not cryptic Elpeddo, new genus
Apical setae on outer ramus of uropod 3 longer than article 2. article 5 of gnathopod 2
cryptic WHdus, new genus
7. Most posterior spines on article 6 of pereopods 1-2 thin, gnathopods stout . Brolgus, new genus
All but one posterior spine on article 6 of pereopods 1-2 thick and short, gnathopods thin
Ganba, new genus
Mandibulophoxus J. L. Barnard
Mandibulophoxus J. L. Barnard, 1957:432.
DIAGNOSIS.?Eyes absent. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae widely spread
towards apex. Article 1 of antenna 2 not ensifonn;
article 3 with 2 setules; facial spines on article 4 in
2 or more rows highly apicad; article 5 thin. Right
mandibular incisor with 5 teeth; molar not tritura-
tive, small, bearing 3 spines, not bearing fuzz;
palpar hump large. Palp of maxilla 1 biarticulate;
inner plate with 4 setae. Setation of maxilla 2 ordi-
nary. Inner plate of maxillipeds ordinary, apex of
palp article 3 not protuberant, dactyl elongate,
apical nail distinct, medium. Gnathopods dissimi-
lar; gnathopod 2 weakly enlarged; article 5 of
gnathopod 1 of ordinary length, free, but cryptic
and short on gnathopod 2, with weak eusirid attach-
ment; palms oblique, hands of gnathopods 1-2
ordinary, ovatorectangular, elongate, poorly setose
anteriorly. Article 6 of pereopods 1-2 weakly setose
posteroproximally. Article 2 of pereopod 3 of
broad form; articles 4-5 of pereopods 3-4 broad;
article 2 of pereopod 3 weakly setose posteriorly.
Pereopod 5 ordinary, article 2 setose ventrally, arti-
cle 3 ordinary, dactyl normal. Epimera 1-2 lacking
long posterior setae, without midfacial setae above
ventral facial ridge; epimeron 3 ordinary. Uroso-
mite 1 generally naked except for sparse ventral
setae; urosomite 3 without dorsal hook or special
process. Peduncle of uropod 1 normally elongate,
without apicoventral spike, with special enlarged
apicolateral spine; peduncular apices of uropods
1-2 not combed; inner ramus of uropod 1 with one
row of marginal spines, some rami continuously
spinose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, elongate article 2 of outer
ramus carrying 3 long apical setae. Telson ordinary,
with 2-4 apical spines on each lobe plus setules.
DESCRIPTION.?Rostrum fully developed. [Fuzz
and calceoli on male antennae unknown.] Prebuc-
cal parts ordinary, poorly separated from each
other, upper lip dominant. Right lacinia mobilis
simple; article I of mandibular palp slightly elon-
gate, palp medium to thick, apex of article 3
oblique, article 2 without outer setae. Lower lip
bearing cones. Outer plate of maxilla 1 with 9
spines, one spine especially thickened. Inner plates
of maxilliped thick, ordinarily setose. Coxae 2-4
without special anterodorsal humps. All posterior
spines on article 6 of pereopods 1-2 thin but stiff,
midapical spine present, enlarged. Article 2 of
pereopod 5 without facial setae. Peduncle of uro-
pod 1 with dorsolateral spines widely spread, me-
dial spines widely spread; peduncle of uropod 2
with medial spines widely spread. Peduncle of uro-
pod 3 lacking extra subapical setae or spines. Tel-
son with midlateral or dorsal setules reduced to one
member on each side.
TYPE-SPECIES.?Mandibulophoxus gilesi J. L.
Barnard, 1957 (monotypy.)
COMPOSITION.?Phoxus uncirostratus Giles, 1890.
RELATIONSHIP. ? Mandibulophoxus resembles
Pontharpinia in the shape of pereopods $-4, noting
especially the apically expanded article 2 of pereo-
pod 3, in the presence of only one midlateral tel-
sonic setule, in the simple right lacinia mobilis (but
flabellate in Pontharpinia), generally in the mandi-
ble and palp, in the short article 2 of antenna 1, the
thin article 5 of antenna 2, in the extra row of ven-
tral spines on article 4 of antenna 2, the elongate
article 2 on the outer ramus of uropod 3, in the
special peduncular spine on uropod 1, and the wide
spread in medial spination on the peduncles of
uropods 1-2.
Mandibulophoxus differs from Pontharpinia in
the simple molar bearing only 2-3 spines, the more
numerous apical spines and absence of a setal brush
on the telson, the reduction in setae on article 2 of
antenna 2, the apical confinement of facial spines
NUMBER 245 91
on article 4 of antenna, 2, the absence of facial setae
on article 2 of pereopod 5, the weakly developed
posteroproximal setation on article 5 of pereopods
1-2, the reduction of setae on the inner plate of
maxilla 1, the absence of posterior setae on article 2
of pereopod 4 and outer setae on article 2 of the
mandibular palp, in the longer nail of the maxil-
lipedal dactyl, in the smaller gnathopod 2, and in
apical details of the second article on the outer
ramus of uropod 3.
REMARKS.?Pontharpinia stimpsoni Stebbing is
removed from this genus to form the type of a new
genus, Basuto, leaving Mandibulophoxus with two
very closely similar species, M. uncirostratus (Giles)
and M. gilesi J. L. Barnard (see Gray and McCain,
1969, for distinctions). The species of Mandibulo-
phoxus are highly advanced tropical and warm-
temperate evolutes, probably with ancestors re-
motely near Pontharpinia. The single species of the
new genus Basuto, from South Africa, forms a rea-
sonably good intergrade between Pontharpinia and
Mandibulophoxus but we do not imply that the
relationship is precise. Basuto has an intermediate
mandibular molar and weak eyes. The cephalic
elongation is approximate to that of Pontharpinia
but the difference between Pontharpinia and Man-
dibulophoxus is not great and Basuto bears the
smoothly rounded posterior lobation on coxa 4 of
Mandibulophoxus, distinct from the situation in
Pontharpinia. Basuto cannot stand on a direct line
to Mandibulophoxus because article 2 of pereopod
3 is narrowed, the outer ramus of uropod 1 is short-
ened, and the dorsal spines are absent. It is there-
fore removed to the Harpiniinae.
Key to the Species of Mandibulophoxus
Article 4 of pereopod 5 about 1.6 times as long as article 5, defining cusp on palms of gnathopods
large and sharp, inner ramus of uropod 3 on female less than half as long as article 1 of
outer ramus M. uncirostratus
Article 4 of pereopod 5 less than 1.3 times as long as article 5, defining cusp on palms of gnatho-
pods small and rounded, inner ramus of uropod 3 on female more than three-fourths as long
as article 1 of outer ramus M. gilesi
Mandibulophoxus uncirostratus (Giles)
Phoxus uncirostratus Giles, 1890:65-66, pi. 2: fig. 2.
Leptophoxus uncirostratus.?Walker, 1904:249.?Nayar, 1959:
14-15, pi. 4: figs. 1-16.
Pontharpinia uncirostrata.? Stebbing, 1906:147.
Mandibulophoxus uncirostratus.?J. L. Barnard, 1957:435-
436.
Pontharpinia uncirostratus.-Pillai, 1957:39-41, fig. 5.
Paraphoxus uncirostrata.?Nayar, 1967:140.
DISTRIBUTION.?Madras coast and Ceylon.
Mandibulophoxus gilesi J. L. Barnard
Mandibulophoxus gilesi J. L. Barnard, 1957:433-435, figs. 1,
2.-Gray and McCain, 1969:189-191, fig. 1.
Mandibulophoxus uncirostratus.?J. L. Barnard, 1960:359
[in part, not Giles]; 1969c: 1% [not Giles].
REMARKS.?Gray and McCain (1969) have dem-
onstrated numerous minute distinctions between
Af. gilesi and M. uncirostratus. To those might be
added the distribution of setae on article 2 of the
mandibular palp. In M. uncirostratus according to
Pillai (1957) the article bears 2 setae in tandem,
whereas according to Barnard (1957) and Gray and
McCain (1969) a pair of contiguous apical setae
occurs on article 2 of the mandibular palp. From
our study of phoxocephalids in Australia we have
learned that this kind of difference has specific
value.
NEW DESCRIPTION.?Facial spine formula on
article 4 of antenna 2 = 2-2-2 or 1-3-2, spines
highly apicad, ventral margin of article 4 with long
blade-like spines in addition to row of long setae;
article 3 of antenna 2 with 2 setae, apicalmost elon-
gate, proximalmost very small. Right mandibular
incisor with 5 teeth, left with 8 sharp teeth in 2
branches; right lacinia mobilis simple, conical,
weakly trifid or irregularly dentate, thin, smaller
than raker spines; left lacinia mobilis flabellate,
with 6-8 teeth; right rakers 8, left 9; each molar a
weak hump, right bearing 3 spines, left 2, spines not
on basal plaque but individually inserted. Pereopods
1-2 with midapical spine on article 6 large, as long
as dactyl or other spines. Apices of uropods 1-2 not
combed.
92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DISTRIBUTION.?Eastern Pacific Ocean from Ya-
quina Bay, Oregon, to southern California, 0-18 m.
Cunmurra, new genus
DIAGNOSIS OF MALE.?Eyes present. [Flagella of
antennae 1-2 probably unreduced in female.] Article
2 of antenna 1 elongate, ventral setae widely spread.
Article 1 of antenna 2 not ensiform; article 3 with
2 setules; facial spines on article 4 in 2 or more
rows; article 5 ordinary in size. Right mandibular
incisor with 3 teeth; molar not triturative, ordinary
to small, pillow-shaped, bearing 4 or more splayed,
semiarticulate spines, usually bearing fuzz; palpar
hump medium. Palp of maxilla 1 biarticulate; in-
ner plate with 4 setae. Setation of maxilla 2 ordi-
nary. Inner plate of maxillipeds ordinary; apex of
palp article 3 not protuberant, dactyl elongate,
apical nail distinct, short to medium. Gnathopods
dissimilar, gnathopod 2 moderately enlarged; arti-
cle 5 of gnathopod 2 of ordinary length, free, elon-
gate on gnathopod 1, with weak eusirid attach-
ment; palms oblique, hands of gnathopods 1-2
elongate and broadened respectively, poorly setose
anteriorly. Article 6 of pereopods 1-2 with postero-
proximal setae. Article 2 of pereopod 3 of broad
form, articles 4-5 of pereopods 3-4 broad to me-
dium, article 2 of pereopods 3-4 not setose poste-
riorly; pereopod 5 ordinary, article 2 naked ven-
trally, article 3 ordinary, dactyl normal. Epimera
1-2 lacking long posterior setae, without midfacial
setae above ventral facial ridge; epimeron 3 ordi-
nary but almost of rounded classification and bear-
ing 2 medium setae. Urosomite 1 generally naked
except for sparse apicoventral setae or spines near
base of uropod 1; urosomite 3 without dorsal hook
or special process. Peduncle of uropod 1 normally
elongate, without apicoventral spike, without special
enlarged apicolateral-medial spine; peduncular
apices of uropods 1-2 not combed; spines if present
on inner ramus of uropod 1 in one row, no rami con-
tinuously spinose to apex, inner ramus of uropod 2
ordinary. Uropod 3 ordinary, article 2 of outer
ramus carrying 2 long apical setae. Telson ordinary,
with only one apical spine on each lobe plus setules.
DESCRIPTION OF MALE.?Rostrum fully developed.
Fuzz on article 1 of antenna 1 in male present,
calceoli on male primary flagellum of antenna 1
present. Calceoli on article 5 of male antenna 2
present, flagellum in male with calceoli. Prebuccal
parts ordinary, poorly separated from each other,
upper lip dominant. Right lacinia mobilis bifid,
thin; mandibular palp thin, article 1 short, article 2
without outer setae, apex of article 3 oblique. Lower
lip bearing cones, outer plate of maxilla 1 with 11
spines, one spine especially thickened. Inner plates
of maxilliped thick, ordinarily setose; outer plates
especially small. Coxae 2-4 without special antero-
dorsal humps. All posterior spines on article 6 of
pereopods 1-2 thick and stiff, midapical seta pres-
ent. Article 2 of pereopod 5 without facial setae.
Peduncle of uropod 1 with dorsolateral spines con-
fined apically, medial spines widely spread; pedun-
cle of uropod 2 with only one medial setule con-
fined apically. Peduncle of uropod 3 lacking extra
subapical setae or spines. Telson with ordinary pair
of midlateral or dorsal setules on each side.
TYPE-SPECIES.?Cunmurra itickerus, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This genus appears to stand near
the ancestral pool of the Brolgus group of genera
in which gnathopod 2 is enlarged and bears a
shortened wrist, and in which the apices of the
peduncles on uropods 1-2 are combed, and in
which epimeron 3 is very simple, poorly setose, pro-
tuberant, and rounded posteroventrally. Cunmurra
is more primitive than other members of the Brol-
gus group because of the presence of 4 spines on
the mandibular molars, the presence of 2 ventral
setae and several posterior setules on epimeron 3,
and the absence of uropodal combs. Gnathopod 2
is weakly enlarged but the wrist is not fully short-
ened. Gnathopod 1 has departed from the hypo-
thetically ideal condition in that it is more slender
than in other members of the group so that Cun-
murra does not form the perfect ancestral grade
visualized for the Brolgus group. Cunmurra retains
the normal birubiin or primitive inner plate on
maxilla 1, which in the Brolgus group often has
reduced numbers and sizes of setae and it also re-
tains the following nonbrolgin features: elongate
article 2 of antenna 1, strongly spinose molar,
stouter spines on article 6 of pereopods 1-2 and
stouter distal articles on pereopods 3-5. Whether or
not Cunmurra retains the elongate article 5 of an-
tenna 2 in the female is unknown because that sex
is undiscovered in Cunmurra.
Cunmurra appears to be in an ancestral grade of
the Birubius mayamyi (11) and B. booleus (15)
groups, especially of the latter, itself a close sibling
NUMBER 245 93
of B. mayamayi. Cunmurra differs from B. booleus
mainly in the distinctive primitive gnathopods and
in the normalcy of the right lacinia mobilis, which
in B. booleus is more complex.
Birubius ularitus (29) is especially close to Cun-
murra itickerus. Besides the slight difference in
gnathopods, Cunmurra itickerus differs from B.
ularitus in the more elongate article 2 of antenna 1,
bifid right lacinia mobilis, smaller outer plate of
the maxilliped, narrower coxa 4, absence of a pos-
terodorsal setule on epiraeron 3, and the absence of
basofacial setae on uropod 1.
Cunmurra must surely find its classificatory home
eventually in the Joubinellinae because of the en-
larged gnathopod 2, elongate article 2 of antenna 1,
and the presence of more than 3 spines on the
mandibular molar. It would be considered to be the
most unspecialized member of that group which is
otherwise composed of aberrant phoxocephalids
having either pelagic and predatorial adaptations
or inquilinous adaptations.
Cunmurra itickerus, new species
FIGURES 30-32
DESCRIPTION OF MALE.?Head about 21 percent
of total body length, greatest width about 63 per-
cent of length; rostrum weakly constricted, broad,
reaching middle of article 2 on antenna 1. Eyes
medium to large, clear of pigment. Article 1 on
peduncle of antenna 1 about 1.2 times as long as
wide, about 2.3 times as wide as article 2, with
medial patch of fuzz, ventral margin with about 13
setules, weakly produced dorsal apex with 4 setules;
article 2 about as long as article 1, with ventral
cycle of 7 setae; primary flagellum with 9 articles,
about 0.66 times as long as peduncle, bearing sev-
eral short aesthetascs; accessory flagellum with 7
articles. Spine formula on article 4 of antenna 2 =
3^t-2, dorsal margin bearing fuzz, ventral margin
with 4 groups of 1?2 short to medium setae, one
ventrodistal long spine; article 5 about as long as
article 4, facial spine formula = 2, dorsal margin
bearing 2 sets of male setae and one calceolus, ven-
tral margin with 2 sets of one seta plus setule, 2
ventrodistal medium spines; flagellum elongate,
flagellar formula = (20-21), 1-4, 6, 8 . . . (18 or 20),
Mandibles with medium pal par hump; right incisor
with 3 teeth; left incisor with 2 teeth and apical
hump; right lacinia mobilis bifid, distal branch much
shorter than proximal, weakly flabellate, narrow,
subbifid, proximal branch simple, pointed, with
marginal denticles; left lacinia mobilis with 5 teeth;
right rakers 7; left rakers 7 plus 2 rudimentaries;
molar in form of bulbous hump, right molar with 6
primarily medium spines, one short spine weakly
disjunct, left molar with 7 primarily medium spines,
one spine weakly disjunct, each molar with plume;
palp article 1 short, article 2 with one long inner
apical seta and 2 other shorter inner setae, article 3
about 1.1 times as long as article 2, oblique apex
with 9 spine-setae, basofacial formula = 0-2. Inner
plate of maxilla 1 ordinary, bearing one long apical
pluseta, one similar apicomedial seta, 2 apicolateral
much shorter setae; palp article 2 with 6 apicalme-
dial marginal spine-setae. Plates of maxilla 2 extend-
ing equally, of similar width, outer with 3 apico-
lateral setae, inner with 3 medial setae. Inner plate
of maxilliped with 2 large thick apical spines, 4
apicofacial setae, one medial seta, outer plate with 6
medial and apical spines, one apicolateral seta; palp
article 1 without apicolateral seta, article 2 with one
apicolateral seta, medial margin of article 2 weakly
setose, article 3 unprotuberant, with 4 facial setae,
one lateral seta, nail of article 4 short, with 2 acces-
sory setules. Coxa 1 not expanded distally, anterior
margin straight; main ventral setae of coxae 1-4 =
5-6-6-3, posteriormost seta of coxae 1-3 shortest;
anterior and posterior margins of coxa 4 parallel,
posterior margin oblique, almost straight, postero-
dorsal corner rounded, posterodorsal margin short,
weakly excavate, width-length ratio of coxa 4 =
2:3. Long posterior setae on article 2 of gnathopods
1-2 and pereopods 1-2 = 3-4-4-4, short anteriors =
2-2-1-1, long anteriors = 1-4-0-0, no others.
Gnathopod 1 with elongate and thin articles 5-6;
gnathopod 2 enlarged, article 5 short, article 6 stout;
width ratios of articles 5-6 on gnathopods 1-2 =
23:33 and 25:43, length ratios = 68:73 and 53:70;
palmar humps ordinary, palms oblique; article 5
of gnathopod 1 elongate, posterior margin almost
flat; article 5 of gnathopod 2 softly triangular. Pereo-
pod 2 stouter than pereopod 1, especially on article 4,
posterior margin of latter more moderately setose,
facial setae formula on article 4 = 3-3, on article
5 = 4-5, main spine of article 5 extending to M. 80
on article 6, article 5 lacking proximoposterior
spines, spine formula of article 6 on both pairs =
4 + 5 plus middistal seta, some spines long; acclivity
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s
FIGURE 30.?Cunmurra itickerus, new species, holotype, male "a," 4.00 mm (Y ? left lacinia
mobilis).
NUMBER 245 95
Gl
FIGURE Sl.?Cunmurra itickerus, new species, holotype, male "a," 4.00 mm.
on inner margin of dactyls of pereopods 1-2 sharp,
produced as tooth, emergent setule long, midfacial
pluseta short. Coxae 5-7 posteroventral setule for-
mula = 1-2-3. Articles 4-5 of pereopods 3-4 of ordi-
nary width, facial spine rows moderately developed,
facial ridge on article 2 of pereopods 3-5 = 1-2-2,
width ratios of articles 2, 4, 5, 6 of pereopod 3 =
48:40:34:16, of pereopod 4 = 69:37:26:11, of pe-
reopod 5 = 88:21:19:8, length ratios of pereopod
3 = 72:30:35:38, of pereopod 4 = 91:55:42:50, of
pereopod 5 = 114:26:26:25, article 2 of pereo-
pod 5 reaching apex of article 4, medial apex of
article 6 bearing 5 digital processes. Posteroventral
corner of epimeron 1 rounded, posterior margin
weakly convex, with setule in notch above corner,
anteroventral margin and face with 2 sets of 3 and
2 setae, posteroventral face with one seta; postero-
ventral corner of epimeron 2 rounded-quadrate, pos-
terior margin almost straight, facial setae = 6 in
tandem; posteroventral corner of epimeron 3
rounded but with weakly protuberant setule sinus,
posterior margin convex, with 3 setule notches, ven-
tral margin with 2 short setae. Urosomite 1 naked,
articulation line complete; urosomites 1-3 unpro-
tuberant. Rami of uropods 1-2 with articulate
enlarged apical nails, outer ramus of uropod 1 with 3
dorsal spines, inner with one, outer ramus of uropod
2 with 2 dorsal spines closely appressed, inner with
none; peduncle of uropod 1 with 2 apicolateral
spines, medially with 4 marginal spines, apicalmost
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
W QPS
FIGURE 32.?Cunmurra itickerus, new species, holotype. male "a," 4.00 mm.
?enlarged; peduncle of uropod 2 with 6-8 dorsal
spines, medially with one tiny apical setule; apicolat-
eral corners of peduncles on uropods 1-2 lacking
comb. Peduncle of uropod 3 with 4 ventral spines,
dorsally with one lateral spine, 2 medial spines; rami
masculine, inner extending to M. 100 on article 1 of
outer: ramus, apex with 2 setae, medial and lateral
margins setose, article 2 of outer ramus short, 0.14,
bearing 2 long setae, apicomedial margin of article 1
with 6 setae, lateral margin with 4 acclivities, spine
formula = 1-1-1-1-1, setal formula = 1-1-1-1-1.
Telson ordinary, length-width ratio = 35:29, al-
most fully cleft, each apex broad, rounded, lateral
acclivity broad, shallow, bearing ordinary lateral
setule, spine next medial longer than setule, mid-
lateral setules ordinary, also bearing rows of male
dorsal denticles. Cuticle with ordinary bulbar set-
ules of varying sizes mixed with pipes, surface bear-
ing extremely fine striations in form of linear fin-
gerprint pattern or simply pebbled, setules espe-
cially short.
ILLUSTRATIONS.?Head reconstructel dorsally, eyes
omitted; lateral view of damaged head partially re-
constructed, with antenna 2 added in disconnected
position.
HOLOTYPE.?AM, male "a," 4.00 mm. Unique.
TYPE LOCALITY.?AM P. 18125, 12 Dec 1939, Kan-
garoo Island, South Australia, light in net at night.
DISTRIBUTION.?South Australia, Kangaroo Island,
neritic.
Brolgus, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae widely spread or
confined apically. Article 1 of antenna 2 not ensi-
form; article 3 with 1-2 setules; facial spines on
article 4 in 2 or more rows; article 5 especially thin
and short. Right mandibular incisor with 3 teeth;
molar not triturative, small, bearing 3 or fewer long
clumped spines with common base, usually not
bearing fuzz; palpar hump small to medium. Palp
of maxilla 1 biarticulate; inner plate with 2 or
fewer setae. Setation of maxilla 2 weak. Inner plate
of maxillipeds ordinary; apex of palp not protuber-
ant, dactyl elongate, apical nail distinct, medium.
Gnathopods dissimilar; hand of gnathopod 2 mod-
erately to strongly enlarged; article 5 of gnathopods
NUMBER 245 97
1-2 of ordinary length to very short, cryptic on
gnathopod 2, with eusirid attachment; palms obli-
que, hand of gnathopod 1 elongate, poorly setose
anteriorly. Article 5 of pereopods 1-2 bearing pos-
teroproximal setae. Article 2 of pereopod 3 of broad
form; articles 4-5 of pereopods 3^1 narrow, article
2 of pereopods 3-4 not setose posteriorly; pereopod
5 ordinary, article 2 naked or weakly setose ven-
trally, article 3 ordinary, dactyl normal. Epimera
1-2 lacking long posterior setae, without midfacial
setae above ventral facial ridge; epimeron 3 of
rounded classification, lacking long setae. Urosomite
1 generally naked except for sparse apicoventral
setae or spines near base of uropod 1; urosomite 3
without dorsal hook or special process. Peduncle of
uropod 1 normally elongate, without apicoventral
spike, with special enlarged apicomedial spine;
peduncular apices of uropods 1-2 combed; spine(s)
on inner ramus of uropod 1 set in one row, no
rami continuously spinose to apex; inner ramus of
uropod 2 ordinary. Uropod 3 ordinary, article 2 of
outer ramus carrying one long apical seta. Telson
elongate, with 1-3 apical spines or setae on each
lobe plus setules.
DESCRIPTION.?Rostrum fully developed, uncon-
stricted. Fuzz on article 1 of antenna 1 in male pres-
ent; calceoli on male primary flagellum of antenna
1 present. Calceoli on article 5 of male antenna 2
present; flagellum in male with calceoli. Prebuccal
parts ordinary, poorly separated from each other,
upper lip dominant. Right lacinia mobilis bifid;
mandibular palp thin, article 1 slightly elongate,
article 2 without outer setae, apex of article 3
oblique. Lower lip bearing cones. Outer plate of
maxilla 1 widi 9?11 spines, one spine especially
thickened. Inner plates of maxilliped thin, poorly
armed. Coxae 2-4 without special anterodorsal
humps. Some posterior spines on article 6 of pereo-
pods 1-2 thin and seta-like, others slightly thicker,
midapical spine or seta absent. Article 2 of pereo-
pod 5 without facial setae. Peduncle of uropod 1
with dorsolateral spines confined apically, medial
spines widely spread; peduncle of uropod 2 with
only one medial spine or setule confined apically.
Peduncle of uropod 3 lacking extra subapical setae
or spines. Telson with ordinary pair of midlateral
or dorsal setules on each side.
TYPE-SPECIES.?Paraphoxus tattersalli J. L. Barn-
ard, 1958 [here designated].
COMPOSITION.?The following new species: Brol-
gus koongarrus, B. mahmak, B. millinus, B. tavelus.
RELATIONSHIP.?Brolgus differs from Kuritus and
Wildus in the presence of only one apical seta on
the outer ramus of uropod 3, and from Ganba in
the presence of several elongate and very thin pos-
terior spines on article 6 of pereopods 1-2 (Ganba
bearing only one such spine, other spines short and
thick), the stouter gnathopod 1, and fully separate
inner plates of the maxillipeds.
The most proximal row of spines on the face of
article 4 on antenna 2 is becoming obsolete in brol-
gins. In that set of armaments we distinguish be-
tween a true spine which lacks plumes and the
plumose setules which approximate the size of the
spines. A spine often carries a subapical trigger.
As shown in the following two keys, two sets of
character alternatives are used to distinguish the
species: the spine and setule formulas on the telson
and the presence or absence of scales on the apical
nails of the rami on uropods 1-2.
Keys to the Species of Brolgus
KEY A
1. Each lobe of telson with 2 stout spines plus 1-2 setules 2
Each lobe of telson with only one stout spine or 1-2 elongate setae plus 1-2 setules 3
2. Rami of uropods 1-2 with apical scale B. koongarrus, new species
Rami of uropods 1-2 lacking apical scale B. miMinus, new species
3. Main element on telson an elongate seta, or exceptionally 2 elongate setal spines, plus 2
setules medial to seta(e) *. tattersaUi
Main element on telson a short, stout spine, plus 1-2 setules either lateral or medial to
spine
4. Rami of uropods 1-2 with apical scale, setules on telsonic lobes lateral to spine
B. tavelus, new species
Rami of uropods 1-2 lacking apical scale, setules on telsonic lobes medial to spine
B. mahmak, new species
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
KEY B
1. Rami of uropods 1-2 lacking apical scale, outer plate of maxilla 1 with 11 spines, inner
ramus of uropod 2 with dorsal spine 2
Rami of uropods 1-2 bearing apical scale, outer plate of maxilla 1 with 9 spines, inner
ramus of uropod 2 naked *
2. Each lobe of telson with 2 stout spines B. millinus, new species
Each lobe of telson with one stout spine or 1-2 elongate setal spines 3
S. Facial ridge on epimeron 2 ordinary, telson with 1-2 elongate setal elements on each lobe
B. tatlmalli
Facial ridge on epimeron 2 obsolescent, very short, telson with only short elements on each
lobe B. mahmak, new species
4. Each lobe of telson with one stout spine and one lateral setule B. tavelus, new species
Each lobe of telson with 2 stout spines separated by setule (or one stout spine and 2 lateral
setules) B. koongarrus, new species
Brolgus tattersalli (J. L. Barnard), new combination
FIGURES SS-36
Parharpinia villosa.?Tattersall, 1922:4-6, pi. 1: figs. 7-14 [not
Haswell, 1879].
Paraphoxus tattersalli J. L. Barnard, 1958:148; 1960:282.
DESCRIPTION OF FEMALE.?Head about 21 percent
of total body length, greatest width almost 70 per-
cent of length; rostrum unconstricted, broad, ex-
ceeding apex of article 2 on antenna 1. Eyes medium,
narrow-reniform, largely occluded with pigment.
Article 1 on peduncle of antenna 1 about 1.75 times
as long as wide, about 1.9 times as wide as article
2, apicoventral margin with about 7 setules, strongly
produced dorsal apex with 3 setules; article 2 about
0.35 times as long as article 1, with apicoventral
rows of 8-9 and 4-6 shorter setae; primary flagellum
with 9-11 articles, about 0.7 times as long as pedun-
cle, bearing aesthetascs on all but first and last
articles; accessory flagellum with 6-9 articles. Spine
formula on article 4 of antenna 2 = 2-1-6-1 or
3-6-1, dorsal margin with notch bearing 3 setal-
spines, ventral margin with 9-10 groups of 1-2 long
to short setae, qne ventrodistal medium spine; arti-
cle 5 about 0.65 times as long as article 4, facial
spine formula = 0, dorsal margin naked, ventral
margin with 5 sets of 1-2 long to short setae, 4 ven-
trodistal long to short spines, 2 of these set as sub-
distal facial spines; flagellum about 0.95 times as
long as articles 4-5 of peduncle combined, with
8-11 articles. Mandibles with medium palpar hump;
right incisor with 3 teeth; left incisor with 3 humps
in 2 branches; right lacinia mobilis bifid, distal
branch much shorter than proximal, flabellate or
simply broad, subbifid or denticulate, proximal
branch simple, pointed, with marginal denticles;
left lacinia mobilis with 5 teeth; right rakers 7-10,
left rakers 8-10, molar in form of short prolusion
or soft cone demarcated mainly by spines, each
molar with 3 long spines; palp article 1 slightly
elongate, article 2 with 1-2 medium inner apical
setae and 3 other shorter inner setae, article 3 about
1.1 times as long as article 2. oblique apex with 8
spine-setae, basofacial formula = 0-1 or 0-2. Inner
plate of maxilla 1 large, broad, bearing one long
apical or facial pluseta, one similar apicomedial
seta; outer plate with 11 spines; palp article 2 with
6 apicalmedial spines and setae. Inner plate of
maxilla 2 shorter and narrower than outer, outer
with 4 apicolateral setae, inner with pair of medial
setae. Inner plate of maxilliped with one large,
thin, feathered apical spine, 2 apicofacial setae, one
medial seta; outer plate with 8-9 medial and apical
spines, no apicolateral setae; palp articles 1-2 naked
laterally, medial margin of article 2 weakly setose,
article 3 with 3 facial setae, one lateral seta; nail of
article 4 long, with one accessory setule. Coxa 1
expanded distally, anterior margin straight; main
ventral setae of coxae 1-4 = 10-9-11?(6?7), poste-
riormost seta of coxae 1-2 slightly shortened; ante-
rior and posterior margins of coxa 4 almost parallel,
posterior margin almost straight, posterodorsal mar-
gin ordinary, V-shaped, width-length ratio of coxa
4=11:12. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 8-8-7-9, long
anteriors = 2-3-0-1, short anteriors = 1-1-1-0.
Gnathopod 1 with elongate hand, gnathopod 2 with
enlarged hand; width ratios of articles 5-6 on
gnathopods 1-2 = 26:36 and 26:55, length ratios =
52:80 and 36:90; palmar humps ordinary, palms
strongly oblique; article 5 of gnathopod 1 ovate,
posterior margin rounded, article 5 of gnathopod 2
NUMBER 245 99
vL
FIGURE 33.?Brolgus tattersalli (J. L. Barnard), female "w," 6.9 mm (v = male "v," 6.7 mm).
100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 34.?Brolgus tattersalli (J. L- Barnard), female "w," 6.9 mm (n - male "n," 5.7 mm;
v = male "v," 6.7 mm).
NUMBER 245 101
FIGURE 35.?Brolgus tattersalli (J. L. Barnard), female "w," 6.9 mm (v = male "v," 6.7 mm;
g = male "g," 6.3 mm; h = female "h," 7.5 mm; a ? female "a," 7.2 mm).
short, triangular, cryptic, almost lobate. Pereopods
1-2 similar, facial setae formula on article 4 = 0
and 0, on article 5 = 0 and 0; main spine of article
5 slender, extending to M. 68 on article 6, article 5
lacking proximoposterior spines; spine formula of
article 6 = 54-5, no middistal seta, some spines
especially thin, acclivity on inner margin of dactyls
of pereopods 1-2 obsolescent, emergent setule al-
most fully immersed, midfacial pluseta ordinary.
Coxae 5-7 posteroventral setule formula = l-(l-S)-
1. Articles 4-5 of pereopods 3-4 narrow, facial spine
rows absent, facial ridge formula on article 2 of
pereopods 3-5 = 0-1-1, width ratios of articles 2, 4,
5, 6 of pereopod 3 = 51:23:20:9, of pereopod 4 =
65:20:14:9, of pereopod 5 = 70:13:10:6, length ra-
tios of pereopod 3 = 63:29:22:33, of pereopod 4 =
80:53:62:78, of pereopod 5 = 89:19:18:24; article
2 of pereopod 5 exceeding middle of article 4;
medial apex of article 6 finely combed, lacking
digital processes. Posteroventral corner of epi-
102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 36.?Brolgus tattersalli (J. L. Barnard) (a ? female "a," 7.2 mm; b - female "b," 6.6
mm; c ? female "c," 6.1 mm; k ? female "k," 6.9 mm; u = male "u," 6.6 mm; v ? male "v,"
6.7 mm; w - female "w," 6.9 mm; z = juvenile "z," 3? mm {abnormal antenna 2)).
meron 1 rounded, posterior margin weakly con-
vex, anteroventral margin with 3 medium setae,
posteroventral face with 3-4 medium setae, setule on
posterodorsal margin; posteroventral corner of
epimeron 2 rounded-quadrate, posterior margin
almost straight, facial setae = 4-6; posteroventral
corner of epimeron 3 rounded, with setule sinus, pos-
terior margin strongly convex, with one setule notch,
ventral margin naked. Urosomite 1 with lateral
setule at base of uropod 1, articulation line com-
plete; urosomite 3 unprotuberant dorsally. Rami of
uropods 1-2 with semi-articulate immersed weak
apical nails lacking scale, outer ramus of uropod 1
with 4-6 dorsal spines, inner with 2-3, outer ramus
of uropod 2 with 3-4 dorsal spines, inner with 1-2
dorsomedial spines; peduncle of uropod 1 with 3-4
apicolateral spines and 2-3 basofacial setae, medially
with 4-5 marginal setae and spines plus apical
NUMBER 245 103
enlarged special spine; peduncle of uropod 2 with
6-7 dorsal spines, medially with one medium apical
spine; apicolateral corners of peduncles on uropods
1-2 with comb. Peduncle of uropod 3 with 6 elon-
gate ventral spines, dorsally with one lateral spine,
one medial spine; rami feminine, inner extending to
M. 85 on article 1 of outer ramus, apex with one
seta, medial and lateral margins naked, article 2 of
outer ramus elongate, 0.30, bearing one long seta,
apicomedial margin of article 1 with one seta,
lateral margin with 5 acclivities, spine formula =
2-2-2-2-2-2, one of each pair elongate, setal for-
mula = 0. Telson elongate, length-width ratio =
7:5, almost fully cleft, each apex narrow, rounded,
lateral acclivity narrow, bearing 1-2 elongate lateral
spine-setae, 2 short medial setules, dorsolateral set-
ules diverse. Cuticle smooth, with fine plumose hairs
scattered on posterior dorsum of body.
OBSERVATIONS (female).?The facial spine formula
on article 4 of female antenna 2 is counted as 1-6-1
but 2 additional long spine-setae occur dorsomedial
to the most distal facial spine and could be counted
in the first group, to wit, 3-6-1; this is true also in
B. millinus and /*. mahmak.
DESCRIPTION OF MALE.?Slighter than female,
with enlarged eyes, shortened peduncle of pleopod
3, weakly crested urosomite 2, heavily setose rami of
uropod 3. Primary flagellum of antenna 1 with 1 +
more articles than in female, articles 1-5 or 1-6
with calceoli and elongated aesthetascs, only distal
articles with elongate aesthetasc, distoventral corner
of article 1 of peduncle with numerous setules.
Articles 3-4 of peduncle on antenna 2 with dorso-
medial fuzz, ventral setae of articles 4-5 fewer and
shorter, dorsal margin of article 5 with 3 sets of
male setae, one spine and 2 calceoli, flagella for-
mula = 38, 1-5, 7, 9 . . . 37. Article 2 of mandibular
palp with 6-8 setae, longer and stouter than in
female; article 3 with basofacial formula of 3-2 or
2-1. Coxae 1-4, gnathopods and pereopods 1-2 with
fewer setae. Spines and setae on article 5 of pereo-
pods 1-2 stouter than in female. Article 2 of pereo-
pod 4 narrower, with spines on anterodistal corner,
anterior marginal setae sparser and shorter, proxi-
mal half of anterior margin concave; pereopod 5
narrower than in female, spines on anterodistal
corners of articles 2-3 shortened. Epimeron 3 less
produced posteriorly, setules smaller. Peduncle of
uropod 1 usually with one additional basofacial
seta, one more medial spine, inner ramus usually
with 2 spines and rudimentary third; peduncle of
uropod 2 with 7-9 spines, outer ramus with 3?4,
inner with 1-2. Rami of uropod 3 masculine.
OBSERVATIONS (male).?Juveniles resembling fe-
males but eyes smaller, becoming sexually differenti-
ated at 5 mm of length. Mature individuals showing
variations, regardless of body size, in numbers of
posterior setae on article 2 of gnathopods, between
4 and 13, usually 9.
VARIATIONS.?Pereopod 4 subject to abnormality
in about 4 percent of specimens with this appendage
intact, abnormal condition reflected in shortness of
article 6, with fewer spines and setae, with occa-
sional shortness of articles 5 and 7, perhaps result-
ing from imperfect regeneration. Juvenile, 3.9 mm,
PPBES 984/1, with right flagellum of antenna 2
composed of only 3 articles, left flagellum with 6
articles.
Variation in telsonic spine patterns is widespread
through all samples; total number of adult speci-
mens closely examined = 606; of these, 435 have the
normal telson with one spine on each lobe; 45 speci-
mens have one spine on one lobe but 2 spines on
the other lobe; 22 specimens have 2 spines on each
lobe; 2 specimens have three spines on one lobe and
2 spines on the other lobe; one specimen has 3
spines on each lobe; finally, one specimen has one
spine on one lobe and no spine on the other lobe.
Specimens with two spines on each telsonic lobe
are not senile though they are generally the largest
individuals of the collections (about 7.0 mm long).
These are also characterized by having more medial
setae on the inner plate of maxilla 1 (Figure 35:
aX2) but no other characteristic has been found to
be correlated with increased telsonic spination, ex-
cept that the distal branch of the right lacinia
mobilis is more elaborate (Figure 36: aMt).
Juveniles of B. millinus and B. mahmak already
have their characteristic telsonic spination in the
size range of 2.4-2.6 mm.
VOUCHER MATERIAL (all illustrated).?CPBS 3IS/
1270: female "w," 6.9 mm; male "v," 6.7 mm.
CPBS 31N/1270: male "n," 5.7 mm. PPBES 920/3:
female "a," 7.2 mm. PPBES 920/4: female "b," 6.6
mm. PPBES 920/2: female "c," 6.1 mm; male "g,"
6.3 mm. PPBES 334: female "k," 6.9 mm. PPBES
984/1: juvenile "z," 3.6 mm. RHM, 22 Oct 1971:
male "u," 6.6 mm. CPBS 31N/77O: female "h," 7.5
mm.
RELATIONSHIP.?Brolgus tattersalli is the type-
104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
species and model for Brolgus and therefore its rela-
tionships are stated in discussions of the other
species of Brolgus.
MATERIAL.?CPBS, 120 samples from SI stations
(546); PPBES, 32 samples from 16 stations (99);
WPBES, 10 samples from 8 stations (12); SBS, 3
samples from 3 stations (4); EBS, 2 samples from 2
stations (2); WAM, one sample (21); AM, 2 samples
(71); RHM, one sample (3).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 0-19 m, sand, silty sand, weed, shell;
off Cape Everard, 77 m. New South Wales: off
Malabar, 49 m, sand, shell; Jervis Bay, 6 m, Halo-
phila, Zostera. South Australia: Kangaroo Island,
neritic. Tasmania: off Tasmanian Coast, 182 m.
Brolgus millinus, new species
FIGURES 37-39
DESCRIPTION OF FEMALE.?Head about 20 percent
of total body length, greatest width about 63 per-
cent of length; rostrum unconstricted, broad, reach-
ing apex of article 2 on antenna 1. Eyes medium,
reniform, clear of pigment. Article 1 on peduncle
of antenna 1 about 1.6 times as long as wide, about
twice as wide as article 2, ventral margin with about
4 apical setules, produced dorsal apex with 2 setules;
article 2 about 0.43 times as long as article 1, with
apicoventral cycle of 6 setae; primary flagellum with
7-10 articles, about as long as peduncle, bearing
several short aesthetascs; accessory flagellum with
5-7 articles. Spine formula on article 4 of antenna
2 = 1?(8?10)?1(+1), dorsal margin with notch bear-
ing 4 setal-spines, ventral margin with 5 groups of
1-2 long to medium setae, one ventrodistal long to
medium seta; article 5 about 0.67 times as long as
article 4, facial spine formula = 0, dorsal margin
naked, ventral margin with 3 sets of 2-3 long to
short setae, 2 ventrodistal medium spines set sub-
distally; flagellum about 1.3 times as long as articles
4-5 of peduncle combined, with 8 articles. Mandi-
bles with medium palpar hump; right incisor with
3 teeth and notch; left incisor with 3 humps in 2
branches; right lacinia mobilis bifid, distal branch
shorter than proximal, broad, subbifid, proximal
branch simple, pointed, with marginal denticles;
left lacinia mobilis with 5 teeth; right rakers 9 plus
1+ rudimentaries; left rakers 10 plus 1+ rudimen-
taries; molar in form of short protrusion demar-
cated mainly by spines, each molar with 3 long
spines, none disjunct; palp article 1 elongate,
article 2 with one medium inner apical seta and
one other shorter inner seta, article 3 about 1.1
times as long as article 2, oblique apex with 5-6
spine-setae, basofacial formula = 0. Inner plate of
maxilla 1 suborbicular, bearing one long apical
pluseta, one similar apicomedial seta; outer plate
with 11 spines; palp article 2 with 2 apical-medial
marginal spines and 3 apicolateral and submarginal
setae. Inner plate of maxilla 2 shorter and narrower
than outer, outer with 3 apicolateral setae, inner
with one medial seta. Inner plate of maxillipcd with
one long, thin apical spine, 2 apicofacial setae,
one medial seta; outer plate with 9 medial and
apical spines, no apicolateral setae; palp articles
1-2 lacking apicolateral seta, medial margin of arti-
cle 2 moderately to weakly setose, article 3 with
3-4 facial setae, one lateral seta, nail of article 4
medium, with one accessory setule. Coxa 1 expanded
distally, anterior margin straight; main ventral
setae of coxae 1-4 = (8-9)-8-8-4. posteriormost
seta of coxae 1-3 slightly shortened; anterior and
posterior margins of coxa 4 almost parallel, poste-
rior margin convex, posterodorsal corner rounded,
posterodorsal margin short, L-shaped, width-length
ratio of coxa 4 = 16:17. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
(2-4)-(2-6)-5-5, long anteriors = 0-2-0-0, short
anteriors = 1-1-1-1, no others. Gnathopods with
enlarged hands, slightly elongate on gnathopod 1;
gnathopod 2 larger than gnathopod 1, with short
and cryptic wrist; width ratios of articles 5-6 on
gnathopods 1-2 = 26:38 and 25:47, length ra-
tios = 55:77 and 42:80; palmar humps ordinary,
palms strongly oblique; article 5 of gnathopod 1
ovate, posterior margin rounded; article 5 of gnatho-
pod 2 short, triangular, almost lobate, cryptic. Pereo-
pods 1-2 similar, facial setae formula on article 4 =
1-1; on article 5 = (0-1) and 1, main spine of article
5 extending to M. 70 on article 6, article 5 lacking
proximoposterior spines; spine formula of article 6
= 4 + 4, no middistal seta, some spines especially
long and thin; acclivity on inner margin of dactyls
of pereopods 1-2 obsolescent, setule almost fully
immersed, midmarginal pluseta ordinary. Coxae 5-7
posteroventral setule formula = 1-1-1; articles 4-5
of pereopods 5?4 narrow, facial spine rows sparse,
facial ridge formula on article 2 of pereopods 3-5 =
0-1-1, width ratios of articles 2, 4, 5, 6 of pereo-
NUMBER 245 105
^ * I
bM
FIGURE 37.?Brolgus millinus, new species, holotype, female "a," 3.9 mm {b = female "b," 45
mm; c = male "c," 4.9 mm).
106 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Gl
FIGURE 38.?Brolgus millinus, new species, holotype, female "a," 3.9 mm (6 ? female "b," 42
mm; c ? male "c," 4.9 mm).
pod 3 = 50:26:21:9, of pereopod 4 = 71:25:15:10,
of pereopod 5 = 78:15:13:7, length ratios of pe-
reopod 3 = 63:30:28:34, of pereopod 4 = 86:59:
70:80, of pereopod 5 = 98:21:20:25; article 2 of
pereopod 5 exceeding middle of article 4; medial
apex of article 6 finely combed, truncate. Postero-
ventral corner of epimeron 1 narrowly rounded,
posterior margin weakly convex, anteroventral mar-
gin with one medium seta, posteroventral margin
with one medium seta; posteroventral corner of
epimeron 2 rounded-quadrate, posterior margin
straight, facial setae = 2, lateral ridge very short to
obsolescent; posteroventral corner of epimeron 3
rounded, weakly protuberant, with setule sinus,
posterior margin convex, with one setule notch,
ventral margin naked. Urosomite 1 with lateral
setule at base of uropod 1, articulation line com-
plete; urosomite 3 unprotuberant dorsally. Rami
of uropods 1-2 with articulate weak apical nails,
lacking scale; outer ramus of uropod 1 with 2-3
dorsal spines, inner with one; outer ramus of uro-
pod 2 with 2 dorsal spines, inner with one dorso-
medial spine; peduncle of uropod 1 with 3 apico-
lateral spines and 2-3 basofacial setae, medially
with 4-5 marginal setae and spines plus special
apical enlarged spine; peduncle of uropod 2 with
NUMBER 245 107
hW
FIGURE 39.?Brolgus millinus, new species, holotype, female "a," 3.9 mm (c ? male "c," 4.9 mm;
h = male "h," 4.6 mm); Brolgus mahmak, new species (g = female "g," 5.25 mm).
5 dorsal spines, medially with one large apical
spine, apicolateral corners of peduncles on uropods
1-2 with comb. Peduncle of uropod 3 with 5?6
ventral spines, some elongate, dorsally with one
lateral spine, one medial spine; rami feminine,
inner extending to M. 60 on article 1 of outer
ramus, apex with one seta, medial and lateral
margins naked, article 2 of outer ramus ordinary,
0.19, bearing one long seta, apicomedial margin of
article 1 with one seta, lateral margin with 3 acclivi-
ties, spine formula = 1-2-2-2, some very elongate.
Telson long, length-width ratio = 28:25, not fully
cleft, each apex narrow, rounded, lateral acclivity
broad, shallow, with elongate lateral and medial
spines separated by ordinary setule, midlateral
setules of equal length. Cuticle with ordinary but
very sparse bulbar setules.
OBSERVATIONS (female).?Spines of outer plate
more elongate than in B. tattersalli, maxillipeds
otherwise like that species; telsonic lobes usually
with 2 (rarely 1 or 3) large spines separated by
setule, medial spine smaller than lateral spine. See
"Observations" (female) under B. tattersalli.
DESCRIPTION OF MALE.?Differing from female in
108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
slightly narrower rostrum, enlarged eyes, elongate
flagellum of antenna 2, broader epimera, narrower
urosome, shortened peduncle of pleopod 3, longer
and more setose rami of uropod 3. Article 1 on
peduncle of antenna 1 with medial patch of fuzz,
ventral margin with numerous setules; each flagel-
lum with additional article, primary flagellum bear-
ing calceoli and elongate aesthetascs. Basal support
articles below head for attachment of antenna 2
much enlarged; articles 3-4 of antenna 2 with dor-
somedial fuzz, article 4 generally with one fewer
facial spine in main row; article 5 lacking both
distofacial spines on article 5, flagellum with up-
wards to 31 articles, flagellar formula = (< 31), 1-4,
6, 8 . . . 30. Distal branch of right lacinia mobilis
more elaborately sculptured than in female, baso-
facial setal formula on article 3 of mandibular
palp = 1-2. Coxae 1-4, gnathopods and pereopods
with fewer setae than in female. Anteroproximal
margin of article 2 on pereopod 4 depressed; pereo-
pod 5 with narrower articles, particularly articles 2
and 6, spines on anterodistal apices of articles 2-3
shorter. Epimera 1-2 broader than in female but
epimeron 3 less deeply produced posteriorly. Uro-
pod 1 usually with one fewer basofacial seta than
in female and one additional dorsal spine, spina-
tion of rami similar to that of female. Telson more
elongate, spines shorter than in female.
ILLUSTRATIONS.?Following items similar to B.
tattersalli and not illustrated: prebuccal mass, upper
lip, outer plate of maxilla 1, maxilliped (but see
outer plate illustrated), pereopodal dactyls. Follow-
ing parts of male similar to those of female and
not illustrated: mandible (but see palp and right
lacinia mobilis), maxillae, maxillipeds, gnathopods,
pereopods 1-4, coxae 1-3. Male and female lower
lips drawn from opposite aspects, that for female
from dorsal aspect, that for male from ventral
aspect.
HOLOTYPE.?NMV, female, 3.9 mm.
TYPE-LOCALITY.?CPBS 23N, 10 Mar 1965, West-
ern Port Bay, Victoria, Australia, 10 m, gravel,
coarse sand, broken shell.
VOUCHER MATERIAL.?CPBS 41N/3: female "b"
4.2 mm (illus). RHM, 22 Oct 1971: male "c," 4.9
mm (illus); male "h," 4.6 mm (illus.). CPBS 32N/
1270: female "k," 4.3 mm.
RELATIONSHIP.?Brolgus millinus differs from B.
koongarrus and B. tavelus in the absence of apical
scales on the rami of uropods 1-2 and in having
11 instead of 9 spines on the outer plate of maxilla
1, and from B. mahmak in the development of 2
full spines on each lobe of the telson and in the
obsolescence or absence of a facial ridge on epi-
meron 2.
Brolgus millinus is a cryptic species very close
to B. tattersalli in which maturation occurs at a
small size, not longer than 5.0 mm, but in which
there is a full development of body armaments
even, in the case of telson and basofacial uropod 1,
a superdevelopment in relation to conditions typi-
cal of B. tattersalli. Brolgus millinus is further dis-
tinguished from B. lattersalli in the short to absent
facial ridge on epimeron 2, the more rectangular
article 1 of antenna 1, the greater height of female
coxa 4. the shorter inner ramus of uropod 3 in the
female, and the shorter article 2 on the outer ramus
of uropod 3.
MATERIAL?CPBS, 71 samples from 21 stations
(159); WPBES, 8 samples from 6 stations (II); SBS,
4 samples from 3 stations (II); RHM, 2 samples (8).
DISTRIBUTION.?Victoria, Western Port, 0.3-19.2
m, fine and coarse sand, muddy sand, sandy gravel,
coarse sand on clay; also nektonic. New South
Wales, off Malabar, 39-66 m.
Brolgus mahmak, new species
FICURES 39 (gT), 4<M2 (part)
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about 70
percent of length; rostrum unconstricted, broad,
reaching or exceeding apex of article 2 on antenna
1. Eyes large, fully black or clear of pigment. Arti-
cle 1 on peduncle of antenna 1 about 1.6-1.7 times
as long as wide, about 1.7 times as wide as article 2,
ventral margin with 4-6 setules, produced dorsal
apex with 1-3 setules; article 2 about 0.35 times as
long as article 1, with apicoventral crescent of 5
setae; primary flagellum with 7-8 articles, about
0.8 times as long as peduncle, bearing several short
aesthetascs; accessory flagellum with 5-6 articles.
Spine formula on article 4 of antenna 2= 3-6-1,
dorsal margin with notch bearing one setule and
3-4 spines, ventral margin with 4-5 groups of 2-3
long to medium setae, 2 ventrodistal long to me-
dium spines; article 5 about 0.7 times as long as
article 4, facial spine formula = 0 or 1, dorsal mar-
gin naked, ventral margin with 2-3 sets of 1-3 long
NUMBER 245 109
FIGURE 40.?Brolgus mahmak, new species, holotype, female "a," 4.54 mm (g ? female "g,"
5^5 mm).
110 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
Fictnx 41.?Brolgus mahmek, new species, holotype, female "a," 4.54 a m .
NUMBER 845 111
mT uT
Fictntz 42.?Upper left: Brolgus mahmak, new spedet, holotype, female "a," 434 mm; lower
Brolgus koongarrus, new species, holotype, male "m," 3.06 mm (m ? holotype; u ?- male V
S.44 mm).
to short setae, 2 ventrodistal medium spines set
somewhat on face; flagellum about 1.0-1.1 times
as long as articles 4-5 of peduncle combined, with
7-8 articles. Mandibles with medium palpar hump;
right incisor with 3 teeth; left incisor with 4 humps
in 2 branches; right lacinia mobilis bifid, distal
branch shorter than proximal, broad, simple, proxi-
mal branch simple, pointed, curved into hook, with
marginal denticles; left ladnia mobilis with 5 teeth
plus 3 accessory teeth; right rakers 5-7 plus 0-2
rudimentaries; left rakers 6-8 plus 1-2 rudimenta-
ries; molar in form of soft cone demarcated mainly
by spines, each molar with 3 long spines; palp arti-
cle 1 slightly elongate, article 2 with 1-2 (rarely 2)
medium apical setae and 1-2 other short inner
setae, article 3 narrow, about 1.1 times as long as
article 2, oblique apex with 5-7 spine-setae, baso-
facial formula = 0. Inner plate of maxilla I broad,
suborbicular, bearing one medium apical pluseta,
one large apicomedial seta; outer plate with 11
spines; palp article 2 with 2 apical-medial margi-
nal spines and 3 submarginal setae. Inner plate of
maxilla 2 shorter and narrower than outer, outer
with 3 apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with small, thin,
unfeathered apical spine, additional thin apicome-
dial spine, 3 apicofacial setae, no medial setae;
outer plate with 7-9 medial and apical spines, no
apicolateral setae; palp articles 1-2 naked laterally,
medial margin of article 2 weakly setose, article 3
with 3-5 facial setae, 2 lateral setae, nail of article
4 short, with one accessory setule. Coxa 1 scarcely
expanded distally, anterior margin straight; main
ventral setae of coxae 1-4 = ($-10)-(S-9)-(7-9)-
(3-5), posteriormost seta of coxae 1-3 slightly short-
ened or seta of coxa 2 scarcely so; anterior and
posterior margins of coxa 4 almost parallel, poste-
rior margin almost straight, posterodorsal corner
narrowly rounded, posterodorsal margin long, al-
most L-shaped; width-length ratio of coxa 4 = 8:9.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = ^ H S - S H ^ M 4 " 6 ) ' sbon
112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
posteriors = 0, long anteriors = 1(4-1 facial)-{l-2
+ 1-2 facials)-O-O, short anteriors = 1-1-1-1, no
others. Gnathopods with enlarged hands, slightly
elongate on gnathopod 1, gnathopod 2 larger than
gnathopod 1; width ratios of articles 5-6 on gnatho-
pods 1-2 = 28:40 and 25=48, length ratios = 53:85
and 37:84; palmar humps small, palms strongly
oblique; article 5 of gnathopod 1 ovate, posterior
margin rounded, short; article 5 of gnathopod 2
short, cryptic, triangular. Pereopods 1-2 similar,
facial setae formula on article 4 = 1 and 1, on arti-
cle 5 = 1 and 2; main spine of article 5 extending
to M. 67 on article 6, article 5 lacking proximo-
posterior spines; spine formula of article 6 = 3 + 4,
no middistal seta; acclivity on inner margin of
dactyls of pereopods 1-2 obsolescent, emergent
setule short, almost fully immersed, midfacial plu-
seta ordinary. Coxae 5-7 posteroventral setule for-
mula = (l-2)-(l-2H- Articles 4-5 of pereopods
3-4 narrow, facial spine rows sparse; facial ridge
formula on article 2 of pereopods 3-5 = 0-1-1;
width ratios of articles 2, 4, 5, 6 of pereopod 3 =
53:26:22:10, of pereopod 4 = 67:25:16:10, of pe-
reopod 5 = 78:15:13:8, length ratios of pereopod
3 = 67:31:30:43, of pereopod 4 = 80:54:65:78, of
pereopod 5 = 95:20:22:27; article 2 of pereopod
5 reaching middle of article 4; medial apex
of article 6 finely combed, undulant. Postero-
ventral corner of epimeron 1 with small protrusion,
posterior margin weakly convex, anteroventral mar-
gin with 1-2 medium setae, posteroventral margin
with 1-2 long setae; posteroventral corner of epi-
meron 2 rounded or weakly protuberant, with weak
sinus, posterior margin weakly convex, facial setae =
1-2; posteroventral corner of epimeron 3 weakly
protuberant, with setule sinus, posterior margin
convex, with one setule notch, ventral margin
naked; epimera 1-2 lacking seta on posterodorsal
margin. Urosomite 1 with lateral setule at base of
uropod 1, articulation line complete; urosomite 3
unprotuberant dorsally. Rami of uropods 1-2 with
articulate apical nails, lacking scale, each with dis-
tinctive jagged flap at base; outer ramus of uropod
1 with 2?3 dorsal spines, inner with 1-2, outer
ramus of uropod 2 with 2 dorsal spines, inner with
1-2 dorsomedial spines; peduncle of uropod 1 with
2 apicolateral spines and 2-4 basofacial setae, me-
dially with 3 marginal setal-spines, plus apical en-
larged special spine; peduncle of uropod 2 with
4-6 dorsal spines, medially with one large apical
spine; apicolateral corners of peduncles on uropods
1-2 with fine comb. Peduncle of uropod 3 with 5-6
ventral spines, dorsally with one lateral spine, one
medial spine; rami feminine, inner extending to
M. 70 on article 1 of outer ramus, apex with one
seta, medial and lateral margins naked, article 2
of outer ramus ordinary, 0.26, bearing one medium
seta, apicomedial margin of article 1 with one seta,
lateral margin with 2-3 acclivities, spine formula =
long and short 2-2-2 or 2-2-2-2, setal formula = 0.
Telson long, length-width ratio = 17:14, almost
fully cleft, each apex narrow, rounded subsharply,
lateral acclivity shallow, weak, bearing short lateral
spine, setule next medial longer than spine, with
occasional second shorter setule, midlateral setules
weakly diverse. Cuticle with scattered ordinary
bulbar setules.
OBSERVATIONS.?See "Observations" under B.
tattersalli.
ILLUSTRATIONS.?Dorsal outline of eyes recon-
structed by approximation on female "g"; antenna
1 shown disarticulate on lateral view of head.
HOLOTYPE.?NMV, female "a," 4.54 mm.
TYPE-LOCALITY.?CPBS 41N/2, Western Port,
Victoria, Australia, 30 Mar 1965, 12.8 m, heavy
reef, large stones, gravel, mud.
VOUCHER MATERIAL.?CPBS 41N/2, female "g,"
5.25 mm (illus.). Male unknown.
RELATIONSHIP.?This species differs from B tat-
tersalli in the presence of a stout and short lateral
spine as the main element on the telsonic lobes and
not an elongate spine-seta as in B. tattersalli. In
B. mahmak the facial ridge on epimeron 2 is
shorter, the inner ramus of uropod 3 on the female
is shorter, the setae on article 2 of antenna 1 are
fewer and more apicad, and the palmar hump on
gnathopod 2 is smaller than in B. tattersalli.
Brolgus mahmak differs from B. millinus in the
failure of the third and medialmost element on
each lobe of the telson to develop fully into a
spine. Characters more minor or difficult to observe
in B. mahmak include the stouter article 2 of an-
tenna 1, the shape of the right lacinia mobilis, the
presence of a short apical spine on the inner plate
of the maxilliped, the presence of a distinctive and
jagged flap at the base of the nails on uropods 1-2
and in the shorter and stouter article 6 and longer
dactyl of pereopod 4, which however are some*
what variable in B. millinus.
NUMBER 245
MATERIAL.?CPBS, 3 samples from 2 stations
(19); WPBES, one sample (SO).
DISTRIBUTION.?Victoria, Western Port, 12.8 m,
heavy reef bottom.
Brolgus tavelus, new species
FIGURES 43, 44
DESCRIPTION OF MALE.?Head about 23 percent
of total body length, greatest width about 70 per-
cent of length; rostrum unconstricted, broad, elon-
gate, evenly tapering, reaching apex of article 2
on antenna 1. Eyes medium, clear of pigment core,
reddish brown. Article 1 on peduncle of antenna 1
about 1.5-1.7 times as long as wide, about 2.S times
as wide as article 2, with medial patch of fuzz,
ventral margin with about 10 setules, weakly pro-
duced dorsal apex with one long seta, one setule;
article 2 about 0.4 times as long as article 1, with
apicoventral crescent of 3-4 setae; primary flagellum
with 6-8 articles, about 0.8 times as long as pe-
duncle, bearing calceolus on articles 1-4, long
aesthetascs on articles 2-(pen)ultimate; accessory
flagellum with 4-6 articles. Spine formula on arti-
cle 4 of antenna 2 = 3-6-1 or 3-5-1, dorsal margin
with notch bearing 3-4 spines, ventral margin with
5-6 groups of one medium or pairs of long to me-
dium setae, 2-3 ventrodistal medium spines, articles
3-4 with dorsomedial fuzz; article 5 about as long
as article 4, facial spine formula = 0-1, dorsal mar-
gin bearing 1-2 calceoli and 3 groups of 3-4 male
Git
FICUMC 43.?-Brolgus tavelus, new species, holotype, male "c," 3J5O mm (y - male "y," 3-63 mm).
114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
FICUKE 44.?Brolgus tavelus, new species, holotype, male "c," 9JSO mm.
setae, ventral margin with one setal-spine pair, sub-
apical facial crescent of 3 medium spines and 2
short setae; flagellum elongate, flagellar formula =
(17-23), (1) 2-4, 6, 8 . . . penultimate. Mandibles
with medium palpar hump; right incisor with 3
teeth and notch; left incisor with 5 humps in 2
branches; right lacinia mobilis bifid, distal branch
much shorter than proximal, subbifid, proximal
branch simple, with marginal denticles; left lacinia
mobilis with 5 teeth plus one accessory tooth; right
and left rakers 4-5 plus 0-2 rudimentaries; molars
demarcated mainly by spines, each molar with 3
long spines; palp article 1 slightly elongate, article
2 with 1-2 long inner apical setae and 1-2 other
short inner setae, article 3 about 0.90 times as long
as article 2, oblique apex with 7-8 spine-setae,
basofacial formula = 0-2. Inner plate of maxilla 1
narrower than in Brolgus mahmak, bearing one
NUMBER MS 115
long apical pluseta, one shorter similar apicomedial
seta; outer plate with only 9 spines; palp article 2
with 2 apical-lateral marginal spines and 3 sub-
marginal setae. Inner plate of maxilla 2 shorter
and narrower than outer, outer with 4 apicolateral
setae, inner with pair of medial setae. Inner plate
of maxilliped with one elongate feathered apical
spine, 3 apicofacial setae, no medial setae; outer
plate with 7-8 medial and apical spine-setae, no
apicolateral setae; palp articles 1-2 naked laterally,
medial margin of article 2 weakly setose, article 3
with 2-3 facial setae, one lateral seta, nail of article
4 long, with one accessory setule. Coxa 1 expanded
distally, anterior margin almost straight; main ven-
tral setae of coxae 1-4 = (8-11 )-(8-10)-9-(5-6),
posteriormost seta of coxae 1-3 slightly shortened;
anterior and posterior margins of coxa 4 almost
parallel, posterior margin slightly convex, postero-
dorsal corner rounded, posterodorsal margin ordi-
nary, V-shaped, width-length ratio of coxa 4 =
14:15 or 10:11. Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 = 2-(0-2)-
2-(2-3), short posteriors = (O-l)-O-O-0, long ante-
riors = (0-l)-(0-l)-O-0, short anteriors = 1-0-1-
(0-1), one long facial on gnathopods 1-2. Gnatho-
pod 1 with elongate hand, gnathopod 2 with en-
larged hand; width ratios of articles 5-6 on gnatho-
pods 1-2 = 28:43 and 26:50, length ratios =
54:97 and 45:102; palmar humps ordinary to
large, palms strongly oblique; article 5 of gnatho-
pod 1 triangular, cryptic, posterior margin short,
article 5 of gnathopod 2 short, triangular. Pereo-
pods 1-2 similar; posterior margin of article 4 on
latter with longer setae, facial setae formula on
article 4 = 0-0 or 1-1, on article 5 = 2-2 and 2-<2-
3), main spine of article 5 extending to M. 90+ on
article 6, article 5 lacking proximoposterior spines,
spine formula of article 6 = 4 + 4, no middistal
seta, some spines especially long; acclivity on inner
margin of dactyls of pereopods 1-2 obsolescent,
emergent setule almost fully immersed, midfadal
pluseta long. Coxae 5-7 posteroventral setule for-
mula = 1-2-1. Articles 4-5 of pereopods 3-4 nar-
row, facial spine rows sparse, facial ridge formula
on article 2 of pereopods 3-5 = 0-1-1; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 50:25:20:10,
of pereopod 4 = 67:23:16:10, of pereopod 5 =
75:13:12:6, length ratios of pereopod 3 = 73:35:
30:38, of pereopod 4 = 100:60:85:92, of pereo-
pod 5 = 115:24:22:29; article 2 of pereopod 5
reaching middle of article 4, medial apex of arti-
cle 6 combed, sigmoid. Posteroventral corner of
epimeron 1 rounded, posterior margin convex, an-
teroventral margin with 1-3 medium setae, postero-
ventral face with one medium seta; posteroventral
corner of epimeron 2 rounded-quadrate, posterior
margin weakly convex, facial ridge extremely weak,
facial setae = 2 crowded forward; posteroventral
corner of epimeron 3 rounded, with setule sinus,
rarely absent, posterior margin strongly convex,
with 0-1 setules, ventral margin naked; epimera
1?2 with small setule on posterodorsal margin set
in weak notch, epimera 2-3 with very short facial
ridge or none. Urosomite 1 with lateral setule at
base of uropod 1, articulation line complete; uro-
somite 3 unprotuberant dorsally. Rarai of uropods
1-2 with articulate apical nails, bearing scale; outer
ramus of uropod 1 with 2-3 dorsal spines, inner
with 1-2, outer ramus of uropod 2 with 1-2 dorsal
spines, inner naked; peduncle of uropod 1 with 2-3
apicolateral spines and one basofacial seta, medi-
ally with 3 marginal spines, plus special apical en*
larged spine; peduncle of uropod 2 with 4-6 dorsal
spines, medially with one large apical spine; apico-
lateral corners of peduncles on uropods 1-2 with
comb, coarse dorsolaterally on uropod 1. Peduncle
of uropod 3 with 6 ventral spines, dorsally with
one lateral spine, one medial spine; rami mascu-
line, inner extending to M. 100 on article 1 of
outer ramus, apex with 2 setae, medial and lateral
margins setose, article 2 of outer ramus ordinary,
0.17, bearing one long seta, apicomedial margin
of article 1 with one spine and one seta, lateral
margin with 3 acclivities, spine formula = 2-2-2-2
or 1-2-2-2, setal formula = 1-1-1-1. Telson elon-
gate, length-width ratio =16:11, not fully cleft,
each apex narrow, rounded, lateral acclivity shal-
low, narrow, bearing ordinary lateral setule, spine
next medial slightly longer or of length equal to
setule, each lobe with row of denticles, midlateral
setules almost alike. Cuticle with sparse, ordinary
bulbar setules.
OBSERVATIONS.?Prebuccal parts moderately ex-
tended forward, epistomal region extended as
rounded lobe from lateral view, epistome and
upper lip fused together but with remnants of
articulation line dorsally, anterior face with cres-
centic rugosity, ventral margin almost truncate,
with tiny midhump (visible if rotated properly);
lower lip as in Brolgus mahmak but mandibular
116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
lobes slightly broader apically; maxilla 1 like that
of B. mahmak (but outer plate with only 9 spines),
shape of inner plate slightly narrower (see illustra-
tion) and palp of different appearance (see illus-
tration), palp with stout apical spine, thinner mar-
ginal spine and S setae; maxilla 2 like that of B.
mahmak; maxilliped generally like that of female
B. mahmak but main apical spine on inner plate
almost twice as long (see illustration), bearing S
apicofacial setae, no ventroapical seta (except for
one attributed as spine), outer plate with 6 spines,
article S of palp with 2 facial setae and one lateral
seta, nail of dactyl (see illustration) much longer
than in B. mahmak; coxae 1-3 as in B. mahmak,
coxa 4 broader, width length ratio averaging
11:12; pereopod 4 about 1.2 times as long as in
B. mahmak relative to pereopods S and 5; apical
nails on rami of uropods 1-2 of this species and
B. koongamts with scale extensions borne on chitin
of ramal body, each scale bearing free setule.
ILLUSTRATIONS.?Calceoli on dorsal margin of
article 5 of antenna 2 not shown; second facial
ridge on article 2 of pereopod 5 apparently arti-
ficial; head of holotype male "c" with eyes in wrong
position owing to preservations! defects, male "y"
with presumed normal eyes and correct insertion
of antenna 1; right mandibular palp of holotype
abnormal in having only one long inner apical seta
on article 2 and no basofacial setae on article 3;
coxa 7 of holotype with setule omitted; postero-
ventral setule sinus on holotype epimeron 3
omitted.
HOLOTYPE.?AM, male "c," 3.50 mm (with right
uropod 3 missing).
TYPE-LOCALITY.?AM P.18125, 12 Dec 1939,
Antechamber Bay, Kangaroo Island, South Aus-
tralia, in net at night.
VOUCHER MATERIAL.?Type-locality, male "y,"
3.63 mm (illus.). Female unknown.
RELATIONSHIP.?The male (female unknown) of
Brolgus tavelus differs from the females of B. tat-
tersalli, B. millinus, and B. mahmak and the males
of B. tattersalli and B. millinus in the telsonic
armament formula, to wit, in B. tavelus the main
apical spine is medial and the setule is lateral,
whereas in the other three species the lateralmost
element is always a spine or elongate seta. The
male of B. koongarrus also has a spine as the lateral
element.
This species and B. koongarrus appear to differ
from the other species in the presence of a scale
adjoining the apical nails on the rami of uropods
1-2. They also have only 9 spines on the outer
plate of maxilla 1 in contrast to the other 3 species
which bear 11 spines.
Together, B. tavelus and B. koongarrus (to fol-
low) differ from B. mahmak in the much narrower
rostrum, the bifid and distally unextended right
lacinia mobilis, unbroadened distal tooth on left
lacinia mobilis, presence in almost all specimens
(except holotypes) of 2 long apical setae on article
2 of mandibular palp, presence of basofacial setae
on article 3 of mandibular palp, absence of a stout
apicomedial spine on inner plate of maxilliped
(apart from normal 3 spine-setae and fine apico-
ventral spine found in all species of Brolgus), much
longer main apical spine on article 5 of pereopods
1-2, longer article 2 of mandibular palp (longer
than article 3), shape of epimeron 3, coarseness of
apicolateral comb on peduncle of uropod 1, the
the uropodal scales, differences in telsonic spina-
tion, and absence of spines on inner ramus of
uropod 2.
MATERIAL.?AM, one sample (8).
DISTRIBUTION.?South Australia, Kangaroo Island,
in net at night.
Brolgus koongarrus, new species
FIGURE 42 (part)
DESCRIPTION OF MALE.?Head about 23 percent
of total body length, greatest width about 22 per-
cent of length; rostrum broad, reaching apex of
article 2 on antenna 1. Eyes subreniform, fairly
large, clear of pigment core, reddish brown. Article
1 on peduncle of antenna 1 about 1.4 times as long
as wide, about 2.25 times as wide as article 2, with
medial patch of fuzz, ventral margin with about
10 setules, weakly produced dorsal apex with 2
setules; article 2 about 0.3 times as long as article
1, with apicoventral cycle of 6 setae; primary flagel-
lum with 7-8 articles, about 0.7 times as long as
peduncle, bearing calceolus on articles 1-4, several
medium aesthetascs; accessory flagellum with 5
articles. Spine formula on article 4 of antenna 2 =
3-5-1 or 3-4-1, dorsal margin with 2 notches bear-
ing 4 and 2 spines, ventral margin with 5-6 me-
dium setae rarely paired with shorter seta, 3 ventro-
distal long and short spines; article 5 about as long
NUMBER MS 117
as article 4, facial spine formula = 0-1, dorsal mar-
gin naked, bearing 3 groups of male setae, occa-
sional calceolus, ventral margin with 2 sets of seta
and spine, subapical facial crescent of 2 spines and
2 short setae; flagellum elongate, flagellar for-
mula = (22-24), 1-8, 5, 7 . . . (21 or 23). Mandibles
with medium palpar hump; right incisor with 3
teeth and notch; left incisor with 5 humps in 2
branches; right lacinia mobilis bifid, distal branch
much shorter than proximal, broad, bifid, branches
gaping, proximal branch simple, pointed, with mar-
ginal denticles; left lacinia mobilis with 5 teeth,
right rakers 4-6 plus 0-1 rudimentary; left rakers
4-5 plus 0-1 rudimentary; molars demarcated
mainly by spines; each molar with S primarily long
spines; palp article 1 slightly elongate, article 2
with 1-3 long inner apical setae and one other
short inner seta, article 3 about 0.9 times as long as
article 2, oblique apex with 6-7 spine-setae, baso-
fadal formula = 0-2 or 1-2. Inner plate of maxilla
1 narrower than in Brolgus mahmak, bearing one
long apical pluseta, one shorter similar apicomedial
seta; outer plate with only 9 spines; palp article 2
with 2 apical-lateral marginal spines and 3 sub-
marginal setae. Inner plate of maxilla 2 shorter and
narrower than outer, outer with 3 apicolateral setae,
inner with one medial seta. Inner plate of maxil-
liped with one thin apical spine, 3 apicofacial setae,
medialmost elongate (contrast B. tavelus), no me-
dial setae; outer plate with 6-8 medial and apical
spines, no apicolateral setae; palp articles 1-2
naked laterally, medial margin of article 2 weakly
setose, article 3 with 3 facial setae, one lateral seta,
nail of article 4 very long (contrast B. tavelus),
with one accessory setule. Coxa 1 expanded distally,
anterior margin almost straight; main ventral setae
of coxae 1-4 = (8-ll)-(8-ll)-(8-9H5-7), posterior-
most seta of coxae 1-3 slightly shortened; anterior
and posterior margins of coxa 4 almost parallel,
posterior margin slightly convex, posterodorsal
corner rounded (but less than B. tavelus), postero-
dorsal margin ordinary, L-shaped, width-length
ratio of coxa 4 = 11:12. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 ?
(l-2>-(l-3)-2-(2-5), short posteriors = 0, long ante-
riors = 1-1-0-0, short anteriors = 0-(0-l)-l-l, 1-2
long facials on gnathopods 1-2. Gnathopod 1 with
elongate hand, gnathopod 2 with enlarged hand;
width ratios of articles 5-6 on gnathopods 1-2 =
28:40 and 26:54, length ratios = 44:104 and
55:109; palmar humps ordinary to large, palms
strongly oblique; article 5 of gnathopod 1 tri-
angular, posterior margin short; article 5 of gnatho-
pod 2 short, cryptic, triangular. Pereopods 1-2
similar; facial setae formula on article 4 = 1 and 1,
on article 5 = 2 and 2, main spine of article 5 extend-
ing to M. 100+ on article 6, article 5 lacking proxi-
moposterior spines, spine formula of article 6 = 3
+ 3, 3 + 4 or 4 + 4, no middistal seta, some spines
especially long; acclivity on inner margin of dactyls
of pereopods 1-2 obsolescent, emergent setule almost
fully immersed, midfacial pluseta long. Coxae 5-7
posteroventral setule formula = 1-2-1. Articles 4-5
of pereopods 3-4 narrow, facial spine rows sparse;
facial ridge formula on article 2 of pereopods 3-5 =
0-1-1; width ratios of articles 2, 4, 5, 6 of pereo-
pod 3 = 55:30:23:11, of pereopod 4 = 70:25:16:10,
of pereopod 5 = 79:15:12:7, length ratios of pe-
reopod 3 = 74:37:32:41, of pereopod 4=111:
70:84:115, of pereopod 5 = 124:27:23:33; article
2 of pereopod 5 almost reaching apex of article
4 or slightly beyond; medial apex of article 6
combed, sigmoid. Posteroventral corner of epimeron
1 rounded, posterior margin convex, anteroventral
margin with 1-3 medium setae, posteroventral face
with one medium seta; posteroventral corner of
epimeron 2 subquadrate, posterior margin weakly
convex, facial ridge short, very weak or absent, facial
setae = one; posteroventral corner of epimeron 3
rounded, posterior margin weakly convex, with 0-2
setule notches, ventral margin naked; epimera 1-2
with small setule on posterodorsal margin set in
weak notch. Urosomite 1 with lateral setule at base
of uropod 1, articulation line complete; urosomite 3
unprotuberant dorsally. Rami of uropods 1-2 with
articulate apical nails, bearing scale, outer ramus
of uropod 1 with 2-3 dorsal spines, inner with 1-2,
outer ramus of uropod 2 with 1-2 dorsal spines,
inner naked; peduncle of uropod 1 with 2-3 apico-
lateral spines and one basofacial seta, medially with
2?3 marginal spines, plus special apical enlarged
spine; peduncle of uropod 2 with 4-5 dorsal spines,
medially with one large apical spine, apicolateral
corners of peduncles on uropods 1-2 with comb,
coarse dorsolaterally on uropod 1. Peduncle of uro-
pod 3 with 6-7 ventral spines, dorsally with one
lateral spine, one medial spine; rami masculine,
inner extending to M. 100 on article 1 of outer
ramus, apex with 2 setae, medial and lateral mar-
gins setose, article 2 of outer ramus ordinary, 0.19,
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
bearing one long seta, medial apex of article 1
with one spine, one seta, lateral margin with 3
acclivities, spine formula = 1-2-2-2 or 2-2-2-2,
setal formula = 1-1-1-1. Telson long, length-width
ratio = 5:4, almost fully cleft, each apex narrow,
rounded, lateral acclivity shallow, narrow, bearing
shorter lateral and longer medial spine separated
by setule longer than lateral spine, lateral spine
occasionally reduced to proportions of setule; each
lobe with row of denticles, midlateral setules almost
alike. Cuticle with sparse ordinary bulbar setules.
OBSERVATIONS.?Prebuccal parts moderately ex-
tended forward, epistomal region extended as
rounded lobe from lateral view, epistome and up-
per lip fused together but with remnants of articu-
lation dorsally, anterior margin with crescentic
rugosity, ventral margin almost truncate, weakly
concave, with tiny midhump (visible if rotated
properly); lower lip generally like that of B. mah-
mak but mandibular lobes broader apically; max-
illa 1 generally like that of B. mahmak, (but outer
plate with only 9 spines), inner plate and palp like
those of B. tavelus; maxilla 2 like that of B. mah-
mak; maxilliped generally like that of B. mahmak
but lacking additional apical spine, bearing 3 apico-
facial setae, no ventrofacial seta (except for one
attributed as spine), outer plate with 6 spines, arti-
cle 3 of palp with 3 facial and one lateral setae,
nail of dactyl much longer than in B. mahmak, like
B. tavelus; coxae 1-3 like female of B. mahmak,
coxa 4 broader, width-length ratio about 11:12, as
in B. tavelus; pereopod 4 about 1.2 times as long as
pereopod 4 of B. mahmak in relation to pereopods
3 and 5; setae on epimeron 1 about 50 percent
longer than in B. tavelus; uropods 1 and 2 with
scales near apical nails as in B. tavelus.
ILLUSTRATIONS.?Mostly omitted because of simi-
larities to B. tavelus and to B. mahmak as detailed
above; seta on medial apex of article 1 of outer
ramus of uropod 3 not shown.
HOLOTYPE.?AM, male "m," 3.06 mm (with right
uropod 3 missing).
TYPE-LOCALITY.?AM P.18125, 12 Dec 1939, An-
techamber Bay, Kangaroo Island, South Australia,
in net at night.
VOUCHER MATERIAL.?Type-locality: male "k,"
3.60 mm; male "u," 3.44 mm (illus.). Female un-
known.
RELATIONSHIP.?The specimens assigned to this
species, like those of Brolgus tavelus, cannot repre-
sent the male of B. mahmak (known only for the
female) because the telson in the majority of these
males bears two fully developed spines on each lobe
and the apical nails on the rami of uropods 1-2
bear scales. Possibly such scales are characters of
males of all these species but until such can be
demonstrated, the feature is considered as a good
taxonomic character. The telson of B. koongarrus
resembles that of B. millinus, hence the best gross
characters of distinction between the species are the
uropodal scales and the 9 spines on the outer plate
of maxilla 1 in B. koongarrus. In B. koongarrus the
apical spine on article 5 of pereopods 1-2 and the
telsonic spines are relatively more elongate than in
B. millinus.
The majority of specimens of B. koongarrus dif-
fers from B. tavelus in the telsonic armament for-
mula. Brolgus koongarrus has 2 thick spines separ-
ated by a setule, whereas B. tavelus has one spine plus
a lateral setule. Occasional specimens of B. koongar-
rus have the thick lateral spine transformed into a
thin seta-spine and often it is rudimentary, thus
approaching the condition in B. tavelus where the
telson has simply one medial spine and one lateral
setule. The aberrant specimens of B. koongarrus
generally have the aberrant condition only on one
side of the telson, the other side being normal.
Brolgus koongarrus also has a broader distal ramus
on the right lacinia mobilis but this feature is so
difficult to observe in proper orientation that we
have not used it as a taxonomic character nor have
we checked it for consistency.
MATERIAL.?AM, one sample (11).
DISTRIBUTION.?South Australia, Kangaroo Is-
land, in net at night.
Elpeddo, new genus
DIAGNOSIS.?Eyes absent. Flagella of antennae
1-2 unreduced in female, article 2 of antenna 1
shortened, ventral setae confined apically. Article
1 of antenna 2 not ensiform; article 3 with 2 set-
ules, facial spines on article 4 in one main row;
article 5 ordinary in size. Right mandibular incisor
with 3 teeth; molar not triturative, medium, bear-
ing 3 long clumped spines with common base, bear-
ing fuzz; palpar hump small to medium. Palp of
maxilla 1 biarticulate, inner plate with 3 setae.
Setation of maxilla 2 ordinary. Inner plate of max-
illipeds ordinary; apex of palp article 3 weakly
NUMBER 245 119
protuberant, dactyl elongate, apical nail distinct,
medium. Gnathopods ordinary, small, similar, gna-
thopod 2 weakly enlarged; article 5 of gnathopods
1-2 of ordinary length, free, without eusirid attach-
ment; palms oblique, hands of gnathopods 1-2
ordinary, ovatorectangular, poorly setose anteriorly.
Article 5 of pereopods 1-2 setose posteroproximally.
Article 2 of pereopod 3 of broad form, weakly
tapering distally; articles 4-5 of pereopods 3-4 nar-
row to medium; article 2 of pereopods 3-4 not
setose posteriorly; pereopod 5 ordinary, article 2
naked ventrally, article 3 ordinary, dactyls normal.
Epimera 1-2 lacking long posterior setae, without
midfacial setae above ventral facial ridge; epimeron
3 of rounded classification and lacking long setae.
Urosomite 1 generally naked except for sparse
apicolateral setae near base of uropod 1; urosomite 3
without dorsal hook or special process. Peduncle of
uropod 1 normally elongate, without apicoventral
spike, with special enlarged apicomedial spine,
peduncle apices of uropods 1-2 combed, spine(s) on
inner ramus of uropod 1 in one row, no rami con-
tinuously spinose to apex, inner ramus of uropod
2 ordinary. Uropod 3 ordinary, elongate article 2
of outer ramus carrying 2 short apical setae. Telson
elongate, with 2 apical spines on each lobe plus
setules.
DESCRIPTION.?Rostrum fully developed. Article
1 of primary flagellum on male antenna 1 elongate,
thickened, densely furnished with aesthetascs, fuzz
absent, calceoli absent. Calceoli on article 5 of male
antenna 2 present; flagellum in male without cal-
ceoli. Prebuccal parts ordinary, poorly separated
from each other, upper lip dominant. Right lacinia
mobilis bifid, flabellate; mandibular palp thin,
article 1 slightly elongate, article 2 without outer
setae, apex of article 3 oblique. Lower lip bearing
cones. Outer plate of maxilla 1 with 11 spines, one
spine especially thickened. Inner plates of maxil-
liped poorly armed, thick. Coxae 2-4 without spe-
cial anterodorsal humps. AH posterior spines on
article 6 of pereopods 1-2 thick and stiff, midapical
spine present. Article 2 of pereopod 5 without facial
setae. Peduncle of uropod 1 with dorsolateral spines
confined apically, medial spines widely spread; pe-
duncle of uropod 2 with only one medial spine con-
fined apically. Peduncle of uropod 3 lacking extra
subapical setae or spines. Telson with ordinary pair
of midlateral or dorsal setules on each side.
TYPE-SPECIES.?Elpeddo kaikai, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This genus differs from Brolgus
and Ganba in the presence of 2 apical setae, albeit
short, on article 2 of the outer ramus on uropod 3,
and from Kuritus in the stout, weakly diverse gna-
thopods with short article 5 on gnathopod 2. El-
peddo appears closest to Wildus, from which El-
peddo differs in the shortness of apical setae on the
outer ramus of uropod 3, in the fully split inner
plates of the maxilliped and in the male by the odd,
lysianassid antenna 1 which in itself may be simply
an aberrancy. The absence of eyes, also a possible
aberrancy, distinguishes Elpeddo from its neigh-
bors.
The presence of double spination on each lobe of
the telson suggests affinities with Brolgus koongar-
rus, B. mahmak, and B. millinus. The two former
species have unserrate right laciniae mobiles whereas
B. millinus has the distal branch serrate as in E.
kaikai. Brolgus millinus has an extremely short
article 2 on the outer ramus of uropod 3, perhaps
one of die best distinguishing characteristics be-
tween the two species.
The unique male is extremely unusual and may
be aberrant; it has the first antenna of a lysianassid
and irregularities in shapes or armaments of uro-
pod 3, urosomite 1, peduncle of uropod 1, antenna
2, and apex of maxillipedal palp, all suggesting
that the first antenna may also be an irregularity,
unique to this specimen and possibly not typical
of the species.
Elpeddo kaikai, new species
FIGURES 45-47
DESCRIPTION OF FEMALE.?Head about 19 per-
cent of total body length, greatest width about 63
percent of length; rostrum unconstricted, not reach-
ing middle of article 2 on antenna 1. Eyes absent.
Article 1 on peduncle of antenna 1 about 1.6 times
as long as wide, about 1.8 times as wide as article
2, ventral margin with about 9 setules, weakly pro-
duced dorsal apex with 2 setules; article 2 about
0.5 times as long as article 1, with apicoventral row
of 6 setae; primary flagellum with 10 articles, about
as long as peduncle, lacking aesthetascs; accessory
flagellum with 7 articles. Spine formula on article
4 of antenna 2 = 7 (or possibly 3-4), dorsal margin
with notch bearing 3 setae and one spine, ventral
120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
yX2 s
FIGURE 45.?Elpeddo kaihai, new species, holotype, female "a," 3.40 mm (y ?? male "y," 3.06 mm).
NUMBER 145 121
G2
FICIIHE 46.?Elpeddo kaikai, new species, holotype, female V 3-*0 mm (y *> male "y," 3.08 mm).
122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
-JJLL
R2
 R1
i7.?Elpeddo.JudkMj,iwm species, holotype, female "a" 3.40 mm ( c - female "c." 3.05
mm; y ? male "y," 3.08 mm).
NUMBER 245 123
margin with 2 groups of 2-3 long to short setae,
one ventrodistal long thin setal-spine; article 5
about 0.7 times as long as article 4, facial spine
formula = 0, dorsal margin naked, ventral margin
with 2 sets of 2 medium to short setae, 3 ventro-
distal short to medium spines, one subdistal facial
spine; flagellum about 1.8 times as long as articles
4?5 of peduncle combined, with 10 articles. Man-
dibles with medium palpar hump; right incisor
with 3 teeth and notch; left incisor with 4 humps
in 2 branches; right lacinia mobilis bifid, distal
branch shorter than proximal, flabellate, denticu-
late, proximal branch weakly flabellate but pointed,
with marginal denticles; left lacinia mobilis with 4
teeth plus 2 accessory teeth; right rakers 6 plus 2
rudimentaries; left rakers 7; molar composed of bulb-
ous hump demarcated mainly by spines, each molar
with 3 long spines, none disjunct, each molar with
plume; palp article 1 slightly elongate, article 2
with one short inner apical seta and one other
shorter inner seta, article 3 about as long as article
2, oblique apex with 6 spine-setae, basofacial for-
mula = 0. Inner plate of maxilla 1 large, bearing one
long apical pluseta, one similar apicomedial seta,
one apicolateral seta; palp article 2 with 6 apical-
medial-lateral marginal spines and setae. Inner
plate of maxilla 2 shorter than outer, outer not
broader than inner, outer with 3 apicolateral setae,
inner with one medial seta. Inner plate of maxil-
liped with one thin apical spine, 3 apicofacial setae,
one medial seta; outer plate with 6 medial and
apical spines, no apicolateral setae; palp article 1
naked, article 2 with one apicolateral seta, medial
margin of article 2 moderately setose, article 3 with
3 facial setae, no lateral setae, nail of article 4
medium, with one accessory setule. Coxa 1 scarcely
expanded distally, anterior margin weakly concave;
main ventral setae of coxae 1-4 = 5-5-5-0, poste-
riormost seta of coxae 1-2 slightly shortened; ante-
rior and posterior margins of coxa 4 almost parallel,
posterior margin convex, posterodorsal corner
sharp-rounded, posterodorsal margin short, V-
shaped, width-length ratio of coxa 4 = 38:45. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = (3-4) A 4-(3-4), short posteriors =
(l-2)-S-(3-5)-4, long anteriors = 1-3-0-0, short
anteriors = 4-3-4-3. Gnathopods weakly enlarged;
wrist of gnathopod 2 short; width ratios of articles
5-6 on gnathopods 1-2 = 27:40 and 28:44, length
ratios = 62:74 and 45:72; palmar humps ordinary,
palms strongly oblique; article 5 of gnathopod I
ovate, posterior margin flat; article 5 of gnathopod
2 short, triangular, almost lobate. Pereopods 1-2
similar; facial setae formula on article 4 = 1 and 2,
on article 5 = 2 and 2, main spine of article 5 ex-
tending to M. 75 on article 6, article 5 lacking
proximoposterior spines; spine formula of article
6 = 2 and 3, no middistal seta, acclivity on inner
margin of dactyls of pereopods 1-2 weak, emergent
setule short, midfacial pluseta short. Coxae 5-7
posteroventral setule formula = 1-1?1. Articles 4?5
of pereopods 3-4 narrow, facial spine rows sparse;
facial ridge formula on article 2 of pereopods 3-5 =
0-1-2 (posterior short); width ratios of articles 2, 4,
5, 6 of pereopod 3 = 44:27:25:11, of pereopod 4 =
68:31:20:9, of pereopod 5 = 90:19:18:8, length
ratios of pereopod 3 = 70:22:30:34, of pereopod
4 = 76:58:37:48, of pereopod 5=110:25:23:27;
article 2 of pereopod 5 almost reaching middle of
article 5; medial apex of article 6 finely combed,
lacking digital processes. Posteroventral corner of
epimeron 1 rounded-quadrate, posterior margin
almost straight, anteroventral margin with 1-3 short
widely spread setae, posteroventral face with 0-1
long seta; posteroventral corner of epimeron 2
rounded-quadrate, posterior margin almost straight,
facial setae = 3; posteroventral corner of epimeron
3 weakly protuberant, with setule sinus, posterior
margin convex, with one setule notch, ventral mar-
gin naked; epimera 1-3 with setule on posterodorsal
margin. Urosomite 1 with lateral setule at base of
uropod 1, articulation line complete. Urosomite 3
unprotuberant dorsally. Rami of uropods 1-2 with
immersed weak apical nails and accessory scale;
outer ramus of uropod 1 with 2 dorsal spines, inner
with one; outer ramus of uropod 2 with 1-2 dorsal
spines, inner with one dorsomedial spine; peduncle
of uropod 1 with one apicolateral spine and one
basofacial seta, medially with 4 marginal spines plus
apical inner enlarged spine; peduncle of uropod 2
with 3-4 dorsal spines, medially with one apical
spine; apicolateral corners of peduncles on uropods
1-2 with comb. Peduncle of uropod 3 with 3 ventral
spines, dorsally with one lateral spine, one medial
spine; rami feminine, inner extending to M. 60 on
article 1 of outer ramus, apex with one seta, medial
and lateral margins naked, article 2 of outer ramus
elongate, 0.50, bearing 2 short setae, medial apex of
article 1 with one seta, lateral margin with 2 acclivi-
ties, spine formula ? 2-2-2, setal formula ? 0.
124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Telson long, length-width ratio ? 6:5, almost fully
cleft, each apex narrow, truncate, lateral acclivity
weak, bearing ordinary lateral setule, spine next
medial of length equal to setule, next spine slightly
shorter, midlateral setules subsequal. Cuticle with
bulbar setules of varying sizes closely packed on
coxae and article 2 of pereopod 5 but rare else-
where, surface bearing fine striations in form of
linear fingerprint pattern, emergent setules espe-
cially short.
OBSERVATIONS (female).?One spine on outer
plate of maxilla 1 in holotype aberrantly trifid.
DESCRIPTION OF MALE (male "y," 3.08 mm, only
specimen known, possibly aberrant).?Antenna 1
with thickened articles 2-3; basal article of flagel-
lum swollen and bearing rows of long aesthetascs
(similar to Lysianassidae). Peduncle of antenna 2
somewhat thinner than in female; anterior triad of
spines and setae on article 4 short, spine formula of
article 4 = 1-7 or 1-6, one long blunt spine on
ventrodistal margin in place of seta found on fe-
male; anterior margin of article 5 with 2 giant cal-
ceoli; right antenna 2 with female-like flagellum
comprising 8 articles about 1.6 times as long as
articles 4?5 of peduncle combined, calceoli absent,
left flagellum masculine but broken after 4 articles,
calceoli absent. Upper lip with small midhump
ventrally, otherwise like female. Right mandible
with 5 rakers; molar and incisor like female, palps
broken and missing, [left mandible lost during dis-
section]. Maxillipedal palp article 4 with apical
nail setule-like and completely immersed, no acces-
sory setules (condition aberrant?). Coxae generally
like those of female, setae identical, coxae 2-3
slightly longer than in female, posterodorsal mar-
gin of coxa 4 even more dorsad than in female.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = (2-3)-3-2-2, short posterior
setae = O-O-l-(2-3), long anterior setae = 1-2-O-0,
short anterior setae = 3 4 4 4, short setae straw-
like. Pereopods 1-2, setae and spine formulas of
article 4 = 1-1, article 5 = 2-2, article 6 = 2 + 3
on both pairs, article 4 thinner than in female, of
pereopod 1 with only 2 groups of 4 and 3 setae pos-
teriorly, of pereopod 2 with groups of 4, 3, and 1.
Article 2 of pereopod 3 slightly thinner than in
female; article 5 of pereopod 5 narrower and more
elongate than in female, medial apex of article 6
on pereopod 5 with comb composed of much en-
larged elements. Shape of epimera generally similar
to that of female, epimeron 1 with 2 anteroventral
setae and one posteroventral seta, none facial [ven-
tral setae probably spread posteriad more than in
female, thus 3 of female spread into groups of 2
and 1]; epimeron 2 with 2 facial setae; epimeron S
lacking dorsal and posterior setules except at corner
sinus. Urosomite 1 with dorsal saddle and invagina-
tion similar to that of lysianassids and ampeliscids.
Medial margin on peduncle of uropod 1 with
spines crowded toward distal margin (see illustra-
tion), basofacial setae = 2 elongate; outer ramus of
uropod 2 with one dorsal spine. Uropod 3 inter-
mediate between masculine and feminine, inner
ramus not elongate but bearing medial setae, outer
ramus with medial setae on article 1. Telson
slightly elongate, bearing irregular basomedial rows
of denticles.
OBSERVATIONS (male).?Possibly aberrant, only
one specimen known.
ILLUSTRATIONS.?Inner plate of maxilliped skewed
laterally in drawing; coxa of pereopod 4 recon-
structed by reference to male "y" and female "c";
illustration of urosome and uropods 1-3 of male
"y" partially reconstructed, very slightly shriveled
on specimen; head of male dorsally similar to
female, not illustrated.
HOLOTYPE?AM, female "a," 3.40 mm.
TYPE-LOCALITY.?SBS C6S3, 22 May 1972, off
Malabar, New South Wales, Australia, 75 m, sandy
gravel.
VOUCHER MATERIAL.?SBS CxS4, male "y," 5.08
mm (illus); SBS C6S2, female "c," 3.05 mm (illus.).
MATERIAL.?SBS, 3 samples (3).
DISTRIBUTION.?New South Wales, off Malabar,
75-92 m, sandy gravel.
Ganba, new genus
DIAGNOSIS.?Eyes present. Flagella of antenna 1-2
unreduced in female. Article 2 of antenna 1 espe-
cially shortened, ventral setae confined apically.
Article 1 of antenna 2 not ensiform, article 3 with
2 setules, facial spines on article 4 in one main row,
article 5 short. Right mandibular incisor with 3
teeth; molar not triturative, small, bearing 3 long
clumped spines with common base, not bearing
fuzz; palpar hump medium. Palp of maxilla 1 bi-
articulate; inner plate with 2 setae. Setation of
maxilla 2 weak. Inner plates of maxillipeds mostly
NUMBER 845 125
fused together; apex of palp article 3 not protuber-
ant, dactyl elongate, apical nail distinct, medium.
Gnathopods dissimilar; gnathopod 2 weakly to
moderately enlarged; article 5 of gnathopod 2 very
short, cryptic, ordinary on gnathopod 1, without
eusirid attachment; palms oblique, hands of gna-
thopods 1-2 elongate, basally broadened, poorly
setose anteriorly. Article 5 of pereopods 1-2 setose
posteroproximally. Article 2 of pereopod 3 of broad
form, articles 4-5 of pereopods 3-4 narrow to me-
dium, article 2 of pereopods 3-4 not setose posteri-
orly; pereopod 5 ordinary, article 2 naked ventrally,
article 3 ordinary, dactyl normal. Epimera 1-2 lack-
ing long posterior setae, without midfacial setae
above ventral facial ridge; epimeron 3 of rounded
classification and lacking long setae. Urosomite 1
generally naked except for sparse apicoventral setae
or spines near base of uropod 1; urosomite 3 without
dorsal hook or special process. Peduncle of uropod
1 normally elongate, without apicoventral spike,
without special enlarged apciolateral-medial spine;
peduncular apices of uropods 1-2 not combed; inner
ramus of uropod 1 naked, no rami continuously
spinose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, article 2 of outer ramus carrying
one long apical seta. Telson ordinary, with only one
apical spine on each lobe plus setules.
DESCRIPTION.?Rostrum fully developed. [Fuzz
on article 1 of antenna 1 in male unknown, calceoli
on male antennae 1-2 unknown.] Prebuccal parts
extended forward, poorly separated from each
other, upper lip dominant. Right lacinia mobilis
bifid, thin; mandibular palp thin, article 1 slightly
elongate, article 2 without outer setae, apex of
article 3 transverse. Lower lip bearing cones. Outer
plate of maxilla 1 with 9 spines, no spine especially
thickened. Inner plates of maxilliped poorly armed,
thick. Coxae 2-4 with special anterodorsal humps.
Some posterior spines on article 6 of pereopods 1-2
thin and seta-like, most thick and stiff, midapical
spine or seta absent. Article 2 of pereopod 5 with-
out facial setae. Peduncle of uropod 1 with dorso-
lateral spines confined apically, medial spines par-
tially spread; peduncle of uropod 2 with only one
medial spine confined apically, Peduncle of uropod
3 lacking extra subapical setae or spines. Telson
with ordinary pair of midlateral or dorsal setules
on each side.
TYPE-SPECIES.?Ganba pellati, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This genus differs from Kuritus
and Wildus in the presence of only one seta on the
outer ramus of uropod 3 and from Brolgus in the
presence of mostly short and stout spines on article
6 of pereopods 1-2, in the much thinner gnathopod
1 and the partially fused inner plates of the maxil-
liped. Unlike other members of the Brolgus group
of genera, Ganba lacks apical combs on the pedun-
cles of uropods 1-2.
Ganba pellati, new species
FIGURES 48, 49
DESCRIPTION OF FEMALE.?Head about 20 percent
of total body length, greatest width about 70 per-
cent of length; rostrum unconstricted, broad, ex-
ceeding apex of article 1 on antenna 1. Eyes me-
dium, largely occluded with pigment. Article 1 on
peduncle of antenna 1 about 1.8 times as long as
wide, about 1.5 times as wide as article 2, ventral
margin with about 6 apical setules, weakly pro-
duced dorsal apex with 2 setules; article 2 about 0.4
times as long as article 1, naked ventrally; primary
flagellum with 8 articles, about 0.8 times as long as
peduncle, bearing numerous short aesthetascs; ac-
cessory flagellum with 5-6 articles. Spine formula
on article 4 of antenna 2 = 2-2-3 or 2-2-4, dorsal
margin with notch bearing one seta and 2 spines,
ventral margin with 3 pairs of long to short setae,
2 ventrodistal medium to short spines; article 5
about 0.67 times as long as article 4, facial spine
formula = 0, dorsal margin naked, ventral margin
with 3 pairs of 2 long to short setae, 3 ventrodistal
short to medium spines, one of these set as subdistal
facial spine; flagellum about as long as articles 4-5
of peduncle combined, with 8 articles. Mandibles
with medium palpar hump; right incisor with 3
teeth and notch; left incisor with 4 humps in 2
branches; right lacinia mobilis bifid, distal branch
much shorter than proximal, forming cusp, proxi-
mal branch simple, pointed; left lacinia mobilis with
4 teeth plus 2 accessory teeth; right rakers 6; left
rakers 6; molar of short protrusion demarcated
mainly by spines, each molar with 3 long spines,
none disjunct; palp article 1 slightly elongate, arti-
cle 2 with one short inner apical seta and one other
short inner seta, article 3 about 1.2 times as long
as article 2, truncate apex with 5-6 spine-setae,
basofacial formula = 0-1. Inner plate of maxilla 1
126 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 4S.?Ganba pellati, new species, holotype, female "a," 5.25 mm (q ?- female "q," 4.45 mm).
NUMBER 245 127
FIGURE 49.?Ganba pellati, new species, holotype, female V 525 mm.
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
especially broad, bearing one long apical pluseta,
one shorter similar apicomedial seta, no apicolateral
setae; palp article 2 with 2 apical-medial marginal
spines and 3-4 submarginal setae. Inner plate of
maxilla 2 shorter and much narrower than outer,
outer with 4 apicolateral setae, inner with one
medial seta. Inner plate of maxilliped with 4 large
thin apical spine-setae, no apicofacial setae, no
medial setae, inner plates mostly fused together;
outer plate with 4 medial and apical spines, no
apicolateral setae; palp article 1 with apicolateral
seta, article 2 with one apicolateral seta, medial
margin of article 2 moderately setose, article S with
4 facial setae, no lateral setae, nail of article 4
medium-short, with one accessory setule. Coxa 1
expanded distally, anterior margin weakly concave,
main ventral setae of coxae 1-4 = 5-5-5-0, all four
with anterodorsal hump, posteriormost seta of coxae
1-3 shortest; anterior and posterior margins of coxa
4 almost parallel, posterior margin almost straight,
posterodorsal corner rounded, posterodorsal margin
short, concave, V-shaped, width-length ratio of coxa
4 = 1:1. Long posterior setae on article 2 of gnatho-
pods 1-2 and pereopods 1-2 = 2-(l-3)-(4-5)-(4-5).
short posteriors = (2-3)-(2-3)-(^*Hs-^)' l o n g an"
teriors = 2-3-0-0, short anteriors = 1-2?2?(1?2), no
others. Gnathopod 1 thin, elongate; gnathopod 2
enlarged, weakly elongate, hands tapering distally;
width ratios of articles 5-6 on gnathopods 1?2 =
23:28 and 26:39, length ratios = 71:100 and 50:88;
palmar humps ordinary, palms strongly oblique;
article 5 of gnathopod 1 elongate, ovate, posterior
margin rounded-truncate; article 5 of gnathopod 2
short, triangular, cryptic. Pereopods 1-2 similar,
facial setae formula on article 4 = (1-2) and (2-3)
on article 5 = (1-2) and (1-3); main spine of article
5 extending to M. 50 on article 6, article 5 lacking
proximoposterior spines; spine formula of article
6 = 2 + 4 plus midposterior seta, spines especially
short, acclivity on inner margin of dactyls of pereo-
pods 1-2 absent, setule fully immersed, midfacial
pluseta short, highly proximal. Coxae 5-7 postero-
ventral setule formula = 1-1-1, articles 4-5 of
pereopods 3-4 medium narrow, facial spine rows
sparse; facial ridge formula on article 2 of pereopods
3-5 = 0-1-1; width ratios of articles 2, 4, 5, 6 of
pereopod 3 = 49:29:25:13, of pereopod 4 = 63:25:
19:12, of pereopod 5 = 79:18:14:9, length ratios
of pereopod 3 = 68:30:30:32, of pereopod 4 =
84:40:38:41, of pereopod 5 = 100:25:22:28; article
2 of pereopod 5 reaching middle of article 4;
medial apex of article 6 finely combed, truncate.
Posteroventral corner of epimeron 1 rounded, pos-
terior margin weakly convex, anteroventral margin
with 4-7 short to medium setae, posteroventral face
with one short seta; posteroventral corner of epime-
ron 2 rounded, weakly protuberant, posterior mar-
gin straight, facial setae = 3-4, posteriormost pair
set vertically if 4 setae present; posteroventral corner
of epimeron 3 weakly protuberant, with setule sinus,
posterior margin weakly convex, with 2 setule
notches, ventral margin naked; epimera 1-2 with
tiny setule on posterodorsal margin. Urosomite 1
with ventral seta and setule at base of uropod 1,
articulation line almost complete; urosomite 3 un-
pro tuber ant dor sally. Rami of uropods 1-2 with
immersed apical nails; outer ramus of uropod 1
with one dorsal spine, inner with none; rami of
uropod 2 naked; peduncle of uropod 1 with S api-
colateral spines and one short basofacial seta, medi-
ally with 3 marginal spines, apicalmost enlarged;
peduncle of uropod 2 with 6-7 dorsal spines, medi-
ally with one large apical spine; apicolateral corners
of peduncles on uropods 1-2 lacking comb. Pedun-
cle of uropod 3 with 5 ventral spines (one disjunct),
dorsally with one lateral spine, one medial spine;
rami feminine, inner extending to M. 66 on article
1 of outer ramus, apex with one seta, medial margin
with one apical seta, article 2 of outer ramus short,
0.14, bearing one long seta, medial margin of article
1 with 3-4 setae, lateral margin with 3 acclivities,
spine formula = 0-1-1-1, setal formula = 1-1?1?1.
Telson ordinary, length-width ratio "-11:10, al-
most fully cleft, each apex narrow, rounded, lateral
acclivity deep, bearing ordinary lateral setule, spine
next medial of length subequal to setule, midlateral
setules diverse. Cuticle with ordinary but sparse
bulbar setules.
OBSERVATIONS.?Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 with occa-
sional short partners or extra short setae between
long members, article 2 of pereopods 1-2 with only
one anterior setule; pereopods 1-2 with one poste-
rior seta on article 6 besides spines; facial setae on
article 5 of pereopods 1-2 highly posteriad.
Young female: Easily recognizable because of gen-
eral specific characters on epimera, head and gnatho-
pods but following external observations included:
article 6 of pereopods 1-2 with only one posterior
spine besides locking spines and one seta; coxae 1-3
NUMBER 245 129
with only S ventral setae each; uropod 1 with one
basofacial seta, 2 apicolateral spines, outer raraus
with one long spine; peduncle of uropod 2 with 4
spines; outer ramus of uropod 3 with one lateral
acclivity bearing one spine only; epimeron 1 with 3
anteroventral and no posterofacial setae, epimeron
2 with 2 setae, epimeron 3 with only 2 posterior
setule notches; article 5 of antenna 2 with only one
posterior set of spine-setae, article 4 like adult.
Juveniles: Smallest 1.47 mm, (Slack-Smith 1),
generally recognizable because of distinct elongation
of article 6 on gnathopod 1; following notes show-
ing reduction in spination on prominent external
appendages: spine formula on article 4 of antenna
2 = 1-1-1 (or 2-1 depending on view); coxae 1-3
with one seta each; epimera 1-2 lacking setae;
peduncle of uropod 1 with one lateral spine, pe-
duncle of uropod 1 with one lateral spine; rami of
uropods 1-2 dorsally naked.
VARIATIONS.?Smaller female "q," 4.45 mm, from
Shepherd 15 (South Australia) better developed
morphologically than holotype female from West-
ern Australia, generally more spinose and setose, up-
per lip with heavy crescentic rugosity, eyes larger
but following parts not compared because of dam-
age to female "q": articles 6-7 of pereopods 1, 2, 4,
5 and uropod 3; holotype with aberrantly vestigial
outer plate of maxilla 2.
ILLUSTRATIONS.?Whole views of gnathopods 1-2
with article 6 not fully flattened, this shape shown
on setoseless outline views; distinctions between
spines and setae on palp of maxilla 1 described
mainly by their positions because these armaments
highly similar to each other; view of left pereopod 3
reconstructed from right appendage; setule on coxa
7 omitted.
HOLOTYPE.?WAM, female "a," 5.25 mm.
TYPE-LOCALITY.?Slack-Smith 1, 4 Dec 1968,
Cheyne Beach, Albany, Western Australia, on weedy
rocks, intertidal.
VOUCHER MATERIAL.?Shepherd 15, female "q,"
4.45 mm (illus.); JLB AUS 13, juvenile "c," 3.34 mm;
/Slack-Smith 1, juveniles, smallest 1.47 mm (6). Male
unknown.
MATERIAL.?Slack-Smith 1, one sample (7); JLB
AUS, one sample (1); Shepherd 15, one sample (1).
DISTRIBUTION.?Albany, Western Australia to
Pearson Island, South Australia, 0-8 m.
Kuritus, new genus
DIAGNOSIS OF MALE.?Eyes present. Flagella of
antennae 1-2 unreduced in male. Article 2 of an-
tenna 1 especially shortened, ventral setae confined
apically. Article 1 of antenna 2 not ensiform, arti-
cle 3 with 2 setules, facial spines on article 4 weakly
in 2 rows, article 5 ordinary in size. Right mandibu-
lar incisor with 3 teeth; molar not triturative, small,
conical, bearing 3 long clumped spines, lacking
fuzz; palpar hump small. Palp of maxilla 1 biartic-
ulate; inner plate with 2 setae. Setation of maxilla
2 weak. Inner plates of maxillipeds ordinary; apex
of palp article 3 not protuberant, dactyl elongate,
apical nail distinct, elongate, Gnathopods small,
similar; article 5 of gnathopods 1-2 of ordinary
length, free, without eusirid attachment; palms
oblique, hands of gnathopods 1-2 ordinary, ovato-
rectangular, elongate, poorly setose anteriorly. Ar-
ticle 5 of pereopods 1-2 setose posteroproximally.
Article 2 of pereopod 3 of broad form, articles 4-5
of pereopods 3-4 narrow to medium, article 2 of
pereopods 3-4 not setose posteriorly; pereopod 5
ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Epimera 1-2 lacking long
posterior setae, without midfacial setae above ven-
tral facial ridge; epimeron 3 of rounded classifica-
tion and lacking long setae. Urosomite 1 generally
naked; urosomite 3 without dorsal hook or special
process. Peduncle of uropod 1 normally elongate,
without apicoventral spike, with special enlarged
apicomedial spine; peduncular apices of uropods
1-2 combed; spine(s) on inner ramus in one row,
no rami continuously spinose to apex, inner ramus
of uropod 2 ordinary. Uropod 3 ordinary, elongate
article 2 of outer ramus carrying 2 medium apical
setae. Telson elongate, with 2 apical spines on each
lobe plus setules, without special dorsal and lateral
spines or setae.
DESCRIPTION OF MALE.?Rostrum fully developed.
Fuzz on article 1 of antenna 1 in male present;
calceoli on male primary flagellum of antenna 1
present. Calceoli on article 5 of male antenna 2
present, flagellum in male with calceoli. Prebuccal
parts extended forward, poorly separated from each
other, upper lip dominant. Right lacinia mobilis
in 2 parts; mandibular palp thin, article 1 short,
article 2 without outer setae, apex of article 3
oblique. Lower lip bearing cones. Outer plate of
maxilla 1 with 9 spines, one spine especially thick-
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ened. Inner plates of maxilliped poorly armed, thick.
Coxae 2-4 without special anterodorsal humps. All
posterior spines on article 6 of pereopods 1-2 thin
and stiff, midapical spine present. Article 2 of
pereopod 5 without facial setae. Peduncle of uropod
1 with dorsolateral spines confined apically, medial
spines widely spread; peduncle of uropod 2 with
only one medial spine confined apically. Peduncle
of uropod 3 lacking extra subapical setae or spines.
Telson with ordinary pair of midlateral or dorsal
setules on each side.
TYPE-SPECIES.?Kuritus nacoomus, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?Kuritus differs from Brolgus and
Ganba in bearing 2 apical setae on the outer ramus
of uropod 3 and from Wildus and Brolgus in the
thin gnathopods that are identical to each other
and on which the wrists are not cryptic. See Para-
phoxus for further relationships.
The type-species has an unusual right lacinia
mobilis in which the two main elements are sepa-
rate, the so-called distal branch being fully articu-
late and the proximal branch therefore resembling
a raker spine. The proximal branch of the right
lacinia mobilis in most phoxocephalids resembles
a raker spine and may actually be a raker spine
fused to the primitive lacinia mobilis. In the case
of Kuritus nacoomus the proximal branch is sepa-
rate but is not counted as a raker spine. In other
respects morphological similarities lie with Brolgus,
the telsonic spination, coxal setation, and epimeral
setation indicating a relationship to B. millinus
and B. mahmak.
Kuritus nacoomus, new species
FIGURES 50, 51
DESCRIPTION OF MALE.?Head about 19 percent
of total body length, greatest width about 60 per-
cent of length; rostrum unconstricted, broad, reach-
ing apex of article 2 on antenna 1. Eyes large,
largely occluded with pigment, ommatidia especially
large. Article 1 on peduncle of antenna 1 about
1.8 times as long as wide, about 2.2 times as wide
as article 2, ventral margin with about 10 setules,
strongly produced dorsal apex with 3 setules;
article 2 about 0.37 times as long as article 1, with
apicoventral cycle of 5 setae; primary flagellum
with 8 articles, about 1.1 times as long as peduncle,
bearing several aesthetascs and calceoli on articles
1-4; accessory flagellum .with 4 articles. Spine
formula on article 4 of antenna 2 = 2-5-1, spines
thin, dorsal margin with 2 notches bearing 3 and 1
spines, ventral margin with 3 groups of 1-3 long to
medium setae, 2 ventrodistal medium and short
spines; article 5 about 0.9 times as long as article 4,
facial spine formula = 1, dorsal margin bearing 2
groups of male setae and 2 calceoli, ventral margin
with 2 pairs of medium and short setae, 2 ventro-
distal medium spines; flagellum elongate, flagellar
formula = 20, 1-3, 5, 7 . . . 19 or 1, 3, 5 Man-
dibles with weak palpar hump; right incisor with 3
teeth, weak notch; left incisor with 2 teeth and
hump; right lacinia mobilis bifid, distal branch
shorter than proximal and fully articulate, broad,
subbifid, proximal branch simple, pointed, with
marginal denticles; left lacinia mobilis with 5 teeth
plus one accessory tooth; right rakers 3; left rakers
4; molars short, conical, demarcated mainly by
spines, each molar with 3 clumped medium spines;
palp article 1 short, article 2 with one long inner
apical seta and one other shorter inner seta, article
3 about 0.95 times as long as article 2, oblique
apex with 7 spine-setae, basofacial formula = 0-3.
Inner plate of maxilla 1 ordinary, bearing one
short stout apical pluseta, one similar apicomedial
seta; denticles on spines of outer plate very thin;
palp article 2 with 2 apical spines and one apico-
medial spine. Plates of maxilla 2 extending sub-
equally, outer slightly broader than inner, outer
with 2 apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with one thin short
apical spine, 2-3 apicofacial setae; outer plate with
4 medial and apical spines, palp articles 1-2 naked
laterally, medial margin of article 2 weakly setose,
article 3 unprotuberant, with 2 facial setae, nail of
article 4 long, with one accessory setule. Coxa 1
scarcely expanded distally, anterior margin almost
straight; main ventral setae of coxae 1-4 = 9-8-8-8,
posterior most seta of coxae 1-3 slightly shortened;
anterior and posterior margins of coxa 4 almost
parallel, posterior convex, posterodorsal corner
rounded, posterodorsal margin long, V-shaped,
width-length ratio of coxa 4 = 18:19. Long poste-
rior setae on article 2 of gnathopods 1-2 and pereo-
pods 1-2 = 2-1-1-2, short anteriors = 2-2-1-1, long
anteriors = 1-1-0-0, no others. Gnathopods slender,
weakly elongate, nearly identical; width ratios of
articles 5-6 on gnathopods 1-2 = 22:29 and 22=28,
NUMBER 245 131
FIGURE 50.?Kuritus nacoomus, new species, holotype, male "a," 4.16 mm.
132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
P5
FIGURE 51.?Kuritus nacoomus, new species, holotype, male "a," 4.16 mm.
NUMBER 245 133
length ratios = 46:74 and 45:74; palmar humps
absent, palms oblique; article 5 of gnathopod 1
ovate, posterior margin rounded-lobate, article 5
of gnathopod 2 similar. Pereopods 1-2 similar;
facial setae formula on article 4 = 1 , on article
5 = 1 ; main spine of article 5 extending to M. 75
on article 6, article 5 lacking proximoposterior
spines; spine formula of article 6 = 3 + 2 plus mid-
distal spine; acclivity on inner margin of dactyls of
pereopods 1-2 obsolescent, emergent setule short,
partly immersed, midfacial pluseta ordinary. Coxae
5-7 posteroventral setule formula = 1-1-1. Articles
4-5 of pereopods 3-4 narrow, facial spine rows
sparse, facial ridge formula on article 2 of pereo-
pods 3-5 = 0-1-1; width ratios of articles 2, 4, 5, 6
of pereopod 3 = 50j29:26:10, of pereopod 4 =
64:22:15:8, of pereopod 5 = 75:14:13:7. length
ratios of pereopod S = 71:32:28:40, of pereopod
4 = 90:54:59:53, of pereopod 5=110:24:23:25;
article 2 of pereopod 5 reaching apex of
article 4; medial apex of article 6 finely combed.
Posteroventral corner of epimeron 1 rounded,
posterior margin straight, anteroventral margin
with one medium seta; posteroventral corner
of epimeron 2 rounded-quadrate, posterior
margin straight, facial setae = 3 in horizontal tan-
dem anteriad, facial ridge absent; posteroventral
corner of epimeron 3 rounded, with setule, poste-
rior margin convex, with one setule, ventral margin
naked. Urosomite 1 with articulation line complete,
urosomite 3 weakly protuberant dorsally. Rami of
uropods 1-2 with articulate apical nails, outer
ramus of uropod 1 with 2 dorsal spines, inner with
one, outer ramus of uropod 2 with one dorsal spine,
inner with one dorsomedial spine; peduncle of
uropod 1 with one apicolateral spine and 4 baso-
facial setae, medially with 5 marginal spines, apical-
most enlarged and set medially; peduncle of uropod
2 with 4 dorsal spines, medially with one apical
spine; apicolateral corners of peduncles on uropods
1-2 with comb, weak on uropod 1. Peduncle of
uropod 3 with 6 ventral spines, dorsally with one
lateral spine, one medial spine; rami submasculine,
inner extending to M. 85 on article 1 of outer
ramus, apex with 2 setae, medial and lateral mar-
gins setose, article 2 of outer ramus elongate, 0.30,
bearing 2 medium setae, apicomedial margin of
article 1 with 2 setules and one seta, lateral margin
with 4 acclivities, spine formula = 1-2-2-2-2, setal
formula = 1-1-1-1-1. Telson long, length-width
ratio = 18:11, not fully deft, each apex narrow,
lateral acclivity weak, with one lateral and one
medial spine separated by setule, midlateral setules
ordinary. Cuticle with ordinary bulbar setules,
emergent setules ordinary, long.
ILLUSTRATIONS.?Lateral view of eyes reconstructed
from damaged condition; dorsal view of head
omitted, like that of male Brolgus millinus but eyes
larger; upper lip shown from oblique ventral view;
apex of right pereopod 2 showing smallest medial
distal spine hidden from lateral view on illustration
of pereopod 1, spine next adjacent considered as
middistal spine hiding distalmost lateral spine.
HoLOTYPE.?WAM male "a," 4.16 mm. Unique.
TYPE-LOCALITY.?WAM 414-73, 2 Sep 1954, Bar-
row Island, Western Australia, presumably surface.
DISTRIBUTION.?Western Australia, Barrow Island,
neritic.
Wildus, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae confined apically.
Article 1 of antenna 2 not ensiform, article 3 with
2 setules, facial spines on article 4 in 2 or more
rows, article 5 especially short. Right mandibular
incisor with 3 teeth; molar not triturative, small,
bearing 3 long clumped spines with common base,
not bearing fuzz; palpar hump medium. Palp of
maxilla 1 biarticulate, inner plate with 2 setae.
Setation of maxilla 2 weak. Inner plates of maxilli-
peds mostly fused together; apex of palp article 3
not protuberant; dactyl elongate, apical nail dis-
tinct, elongate. Gnathopods scarcely dissimilar,
gnathopod 2 weakly enlarged; article 5 of gnatho-
pod 2 very short, cryptic, slightly more elongate
on gnathopod 1, without eusirid attachment; palms
oblique, hands of gnathopods 1-2 rectangular,
slightly elongate, poorly setose anteriorly. Article 5
of pereopods 1-2 setose posteroproximally. Article
2 of pereopod 3 of broad form, articles 4-5 of pe-
reopods 3?4 narrow to medium, article 2 of pereo-
pods 3-4 not setose posteriorly; pereopod 5 ordi-
nary, article 2 naked ventrally, article 3 ordinary,
dactyl normal. Epimera 1-2 lacking long posterior
setae, without midfacial setae above ventral facial
ridge; epimeron 3 of rounded classification and
lacking long setae. Urosomite 1 generally naked;
134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
urosomite 5 without dorsal hook or special process.
Peduncle of uropod 1 normally elongate, without
apicoventral spike, with special enlarged apicome-
dial spine; peduncular apices of uropods 1-2
combed; spine(s) on inner ramus of uropod 1 in
one row, no rami continuously spinose to apex,
inner ramus of uropod 2 ordinary. Uropod 3 ordi-
nary, article 2 of outer ramus carrying 2 medium
to long apical setae. Telson ordinary to elongate,
with only one apical spine on each lobe plus
setules.
DESCRIPTION.?Rostrum fully developed. Fuzz on
article 1 of antenna 1 in male present; calceoli on
male primary flagellum of antenna 1 present. Cal-
ceoli on article 5 of male antenna 2 present; flagel-
lum in male with calceoli. Prebuccal parts ordi-
nary, poorly separated from each other, upper lip
dominant. Right lacinia mobilis bifid, thin; man-
dibular palp thin, article 1 slightly elongate, article
2 without outer setae, apex of article 3 almost
transverse. Lower lip bearing cones. Outer plate of
maxilla 1 with 9 spines, no spine especially thick-
ened. Inner plates of maxilliped thin, poorly
armed. Coxae 2-4 without distinct anterodorsal
humps. $ome posterior spines on article 6 of pereo-
pods 1-2 thin and seta-like, others thick and stiff,
midapical spine or seta absent. Article 2 of pereo-
pod 5 without facial setae. Peduncle of uropod 1
with dorsolateral spines confined apically, medial
spines widely spread, peduncle of uropod 2 with
only one medial spine confined apically. Peduncle
of uropod 3 lacking extra subapical setae or spines.
Telson with ordinary pair of midlateral or dorsal
setules on each side.
TYPE-SPECIES.?Wildus thambaroo, new species.
COMPOSITION.?Wildus mullokus, new species;
Parharpinia fuegiensis Schellenberg, 1931; Para-
phoxus waipiro J. L. Barnard, 1972b.
RELATIONSHIP.?Wildus differs from Brolgus and
Ganba in bearing 2 setae on the outer ramus of
uropod 3 and differs from Kurilus in the stouter
and more diverse gnathopods with cryptic article 5
on gnathopod 2. See Elpeddo for further differ-
entiation.
Key to the Species of Wildus
1. Epimeron S with S ventral setae W.t fueglentis, new combination
Epimeron 3 lacking ventral setae 2
2. Inner rami of uropods 1-2 dorsally naked, peduncle of uropod 2 with 2 apicolateral spines
WJ waipiro, new combination
Inner rami of uropods 1-2 with one dorsal spine each, peduncle of uropod 2 with one api-
colateral spine S
3. Coxa 4 with long setae, article 6 of pereopods 1-2 with one elongate setal spine
W. mullokus, new species
Coxa ? lacking long setae, article 6 of pereopods 1-2 with three elongate setal spines
W. thambaroo, new species
Wildus thambaroo, new species
FIGURES 52-54
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about
60 percent of length; rostrum unconstricted, broad,
short, reaching middle of article 2 on antenna 1.
Eyes medium, largely occluded with pigment. Arti-
cle 1 on peduncle of antenna 1 about 1.4 times as
long as wide, about 1.7 times as wide as article 2,
ventral margin with about 4 apical setules, weakly
produced dorsal apex with one setule; article 2
about 0.5 times as long as article 1, with apico-
ventral cycle of 4-5 setae; primary flagellum with
6 articles, about 0.75 times as long as peduncle,
bearing aesthetascs; accessory flagellum with 4 arti-
cles. Spine formula on article 4 of antenna 2 =
3-3-2-1, dorsal margin with notch bearing seta
and 2 spines, ventral margin with 3 groups of 1-2
long to medium setae, one ventrodistal long spine;
article 5 about 0.65 times as long as article 4, facial
spine formula = 0, dorsal margin naked, ventral
margin with 2 pairs of long to short setae, 4 ventro-
distal short to medium spines, 2 of these set as
subdistal facial spines; flagellum 1.1-1.25 times as
long as articles 4-5 of peduncle combined, with
6-7 articles. Mandibles with medium palpar hump;
right incisor with 3 teeth and notch; left incisor
with notch and 4 humps in 2 branches; right la-
cinia mobilis bifid, distal branch much shorter than
NUMBER 245 155
FIGURE 52.?Wildus thambaroo, new species, holotype, female "a," 2.43 mm (q - male "q." 2.56
mm; to ? female "w," 2.47 mm).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 53.?Wildus thambaroo, new species, holotype, female "a," 2-43 mm (q ? male "q," 2.36
mm; w ? female "w," 2.47 mm).
NUMBER 245 137
FicimE. 54.?Wildus thambaroo, new species, holotype, female
"a," 2.43 mm.
proximal, narrow, simple, proximal branch simple,
pointed, with marginal denticles; left lacinia mo-
bilis with 5 teeth plus accessory hump; right rakers
4; left rakers 4; molar in form of short protrusion
demarcated mainly by spines, right molar with 3
long spines, none disjunct; left molar with 3 long
spines, none disjunct; palp article 1 slightly elon-
gate, article 2 with one medium inner apical seta
and one other shorter inner seta, article 3 about
as long as article 2, weakly oblique apex with 5
spine-setae, basofacial formula = 0. Inner plate of
maxilla 1 large, bearing one apical pluseta, one
longer apicomedial seta; palp article 2 with 5
apical-lateral-medial marginal spines and setae.
Inner plate of maxilla 2 shorter and slightly nar-
rower than outer, outer with 3 apicolateral setae,
inner with one medial seta. Inner plates of max-
illiped partly fused, each with one large, thin apical
spine-seta, 3 apicofacial setae, no medial setae;
outer plate with 4 medial and apical spines, no
apicolateral setae; palp articles 1-2 laterally naked,
medial margin of article 2 weakly setose, article 3
with 2 facial setae, no lateral setae, nail of article
4 medium, with one accessory setule. Coxa 1 ex-
panded distally, anterior margin straight; main
ventral setae of coxae 1-4 = 4 5 4-0, posteriormost
seta of coxae 1-3 slightly shortened; anterior and
posterior margins of coxa 4 almost parallel, poste-
rior margin convex, posterodorsal corner rounded,
posterodorsal margin short, V-shaped, width-length
ratio of coxa 4 ? 8:9. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
2-(2-3)-<1-SH4-5)' s h ? r t posteriors = 1-0-0-0, long
anteriors = 1-2-0-0, short an tenors = 2-2-1-1, no
others. Gnathopods weakly enlarged, gnathopod 2
larger, wrists very short; width ratios of articles
5-6 on gnathopods 1-2 = 24:36 and 21:40, length
ratios = 44:72 and 35:72; palmar humps ordi-
nary, palms strongly oblique; article 5 of gnatho-
pod 1 triangular, posterior margin rounded, lobate;
article 5 of gnathopod 2 short, triangular. Pereo-
pods 1-2 similar, facial setae formula on article
4 = 0 and 1, on article 5 = 1 and 2; main spine of
article 5 extending to M. 67 on article 6, article 5
lacking proximoposterior spines; spine formula of
article 6 = 4 + 3 and no middistal seta, some spines
especially thin; acclivity on inner margin of dactyls
of pereopods 1-2 obsolescent, emergent setule long,
midfacial pluseta ordinary, highly proximal. Coxae
5-7 posteroventral setule formula = 1-2-1; articles
4?5 of pereopods 3-4 narrow, facial spine rows
sparse, facial ridge formula on article 2 of pereo-
pods 3-5 = 0-1-1; width ratios of articles 2, 4, 5, 6
of pereopod 3 = 55:27:24:11, of pereopod 4 =
76:26:19:11, of pereopod 5 = 82:16:14:9, length
ratios of pereopod 3 = 75:25:29:34, of pereopod
4 = 90:48:40:49, of pereopod 5 = 103:20:22:28;
article 2 of pereopod 5 reaching middle of article
4; medial apex of article 6 finely combed and trun-
cate. Posteroventral corner of epimeron 1 narrowly
rounded, posterior margin weakly convex, antero-
ventral margin with one short seta, posteroventral
face lacking setae; posteroventral corner of epi-
meron 2 rounded, posterior margin weakly convex,
facial setae = 2; posteroventral corner of epimeron
3 weakly protuberant, with setule sinus, posterior
margin weakly convex, with one setule notch, ven-
tral margin naked. Urosomite 1 naked, articula-
tion line almost complete; urosomite 3 unprotuber-
ant dorsally. Rami of uropods 1-2 with articulate
enlarged apical nails, outer rami of uropods 1-2
and inner of uropod 1 with one accessory setule,
outer ramus of uropod 1 with one dorsal spine,
inner with one, outer ramus of uropod 2 with one
dorsal spine, inner with one dorsomedial spine;
138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
peduncle of uropod 1 with one apicolateral spine
and one basofacial seta, medially with 3 marginal
spines, apicalmost enlarged, plus one larger spine
at base of inner ramus; peduncle of uropod 2 with
2 dorsal spines, medially with one small apical
spine; apicolateral corners of peduncles on uro-
pods 1-2 with comb. Peduncle of uropod 3 with 5
ventral spines, one of these elongate, dorsally with
one lateral spine, one medial spine; rami feminine,
inner extending to M. 45 on article 1 of outer
ramus, apex with one seta, medial and lateral mar-
gins naked, article 2 of outer ramus elongate, 0.34,
bearing 2 medium and long setae, apicomedial mar-
gin of article 1 with one seta, lateral margin with
3 acclivities, spine formula = 2-2-2-2, setal for-
mula ?? 0. Telson weakly elongate, length-width
ratio = 12:11, not fully cleft, each apex narrow,
rounded-truncate, lateral acclivity broad, shallow,
bearing ordinary lateral setule, spine next medial
longer than setule, midlateral setules subequal in
size. Cuticle with ordinary but sparse bulbar setules.
DESCRIPTION OF MALE.?Immature, similar to
female but eyes larger, rostrum slightly constricted
subapicad (dorsal view). Antenna 1 like female, no
medial fuzz on article 1. Antenna 2 scarcely elon-
gate, flagellum 9-articulate, spine formula on arti-
cle 4 = 3-3-2-1, on article 5 = (1 tiny setule). Right
mandible with 3 rakers and one rudimentary;
molar with 2 spines and one rudimentary; palp
with only 4 apical setae on article 3. Inner plate
of maxilliped with only 2 dorsofadal setae, palp
article 3 with only one facial seta. Coxa 4 larger
than in female but of similar shape. Article 2 of
gnathopods 1-2 each with 2 long posterior setae;
article 6 of these gnathopods slightly thinner than
in female. Article 2 of pereopods 1-2 with 2 long
posterior setae each, facial setal formulas on article
4 of pereopods 1-2 = 0-0, on article 5 = 2-2 (1
long, 1 short), on article 6 = (3-1) and (3-1), with
one seta next to each of 2 proximal spines in the
row of 3. Article 2 of pereopod 3 narrower than in
female (see illustration); pereopod 5 relatively
larger than in female but shape of article 2 gener-
ally similar. Uropod 3 with inner ramus elongate,
reaching to apex of article 1 on outer ramus, naked
except for 2 apical setae. Telson elongate, length-
width ratio = 16 : 11.
DESCRIPTION OF JUVENILE (smallest 1.36-1.40
mm).?Eye very small. Spine formula on article 4
of antenna 2 = 2-3-1. Setae of coxae 1-3 = 3-3-3.
Facial setae on article 5 of pereopods 1-2 = 1-2;
spines on article 6 = 2 + 2 and 2 + 2. Articles 4-6
of pereopod 3 lacking posterior spines and setae.
Epimera 1-2 lacking setae. Peduncle of uropod 1
with one apicolateral spine; rami of uropods 1-2
naked or outer ramus with one dorsal spine; pe-
duncle of uropod 2 with 1-2 apicolateral spines.
Outer ramus of uropod 3 with 0-1 lateral acclivity
and 0-1 lateral spine, article 2 of outer ramus
relatively longer than in adult.
ILLUSTRATIONS.?Article 3 of maxillipedal palp
not flattened; pereopod 4 illustrated in 2 sections
joined together, articles \-4 from left side, articles
5-7 from right side (medial view on latter portion);
uropod 3 of female holotype aberrant (Figure 53:
R3), bearing excessive total of 2 dorsolateral spines
and no elongate ventral spine, one seta on article 2
of outer ramus missing; see normal peduncle of
right uropod 3 in Figure 53: nR3t.
HOLOTYPE.?WAM, female "a," 2.43 mm.
TYPE-LOCALITY.?JLB AUS 13. 30 Sep 1968,
Middleton Beach, Albany, Western Australia, inter-
tidal wash of sandy rocks, coralline algae.
VOUCHER MATERIAL.?Type-locality: male "q,"
2.36 mm (illus.); juvenile "c," 1.40 mm; female "w,"
2.47 mm (illus.). Slack-Smith 1: juvenile "j," 1.36
mm.
RELATIONSHIP.?See Wildus mullokus.
MATERIAL.?JLB AUS. 3 samples (14); Slack-
Smith 1, one sample (7).
DISTRIBUTION.?Western Australia, Albany, inter-
tidal.
Wildus mullokus, new species
FIGURES 55-57
DESCRIPTION OF MALE.?Head about 19 percent
of total body length, greatest width about 60 per-
cent of length; rostrum unconstricted, broad,
reaching middle of article 2 on antenna 1. Eyes
medium, clear of pigment core, but stained eosin
in preservative. Article 1 on peduncle of antenna 1
about 1.4 times as long as wide, about twice as wide
as article 2, apicoventral margin with about 9
setules, strongly produced dorsal apex with 3
setules; article 2 about 0.45 times as long as article
1, with apicoventral cycle of 5 setae; primary flagel-
lum with 7 articles, about 0.85 times as long as
peduncle, bearing several aesthetascs and one
NUMBER 245 139
calceolus each on articles 1-3; accessory flagellum
with 5 articles. Spine formula on article 4 of an-
tenna 2 = 1-5-1, dorsal margin with notch bear-
ing 2 spines, and proximally another spine, ventral
margin with 3 groups of 1-3 short-medium setae,
2 ventrodistal long to medium spines; article 5
about 0.8 times as long as article 4, facial spine
formula = 1, dorsal margin bearing 2 calceoli and
3 groups of male setae, ventral margin with 1-2
sets of 1-2 setae, ventrodistally with 2 medium
spines and 2 setules; flagellum elongate, (lagcllar
formula = 21. 1-3. 5. 7 . . . 21 or 1. 2. 4, 6 . . . 20.
Mandibles with weak palpar hump; right incisor
with 3 teeth and midnotch; left incisor with 5
humps in 2 branches; right lacinia mobilis bifid,
distal branch little shorter than proximal, distal
branch narrow, weakly subbifid, proximal branch
simple, weakly pointed; left lacinia mobilis with
5 teeth and 2 accessory teeth; right rakers 5; left
rakers 6 plus one rudimentary; molar in form of
short protrusion crowned with 3 long spines; palp
article 1 slightly elongate, article 2 with one long
inner apical seta and one other shorter inner seta,
article 3 about as long as article 2, weakly oblique
apex with 6 spine-setae, basofacial formula = 0-2.
Inner plate of maxilla 1 ordinary, bearing one
medium long apical pluseta, one similar apico-
medial seta, no apicolateral setae; palp article 2
with one apical spine on protrusion, 2 apicolateral
spine-setae, one apicomedial seta and one facial
seta. Inner plate of maxilla 2 shorter than outer,
outer plates slightly broader than inner, outer with
3 apicolateral setae, inner with one medial seta.
Inner plate of maxilliped with 3 large apical spine-
setae, one medial seta, one small apicomedial seta,
plates fused medially more than normally; outer
plate with 5 medial and apical spines, no apico-
lateral setae; palp articles 1-2 without apicolateral
setae, medial margin of article 2 weakly setose,
article 3 with 2 facial setae, no lateral setae, nail of
article 4 long, with one accessory setule. Coxa 1
scarcely expanded distally, anterior margin almost
straight; main ventral setae of coxae 1-4 = 6-7-6-2,
posteriormost seta of coxae 1-3 slightly shortened;
anterior and posterior margins of coxa 4 almost
parallel, posterior margin convex, posterodorsal
corner rounded, posterodorsal margin ordinary, V-
shaped, width-length ratio of coxa 4 = 5:6. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 1-2-2-3, long anteriors = 0-1-0-0,
short anteriors = 2-2-1-1. Gnathopods generally
ordinary, subequal in size; width ratios of articles
5-6 on gnathopods 1-2 = 24:40 and 25:41, length
ratios = 55:88 and 47:80; palmar humps ordinary,
palms strongly oblique; article 5 of gnathopod 1
with posterior margin rounded; article 5 of gnatho-
pod 2 with posterior margin triangular. Pereopods
1-2 similar, posterior margin of article 4 moder-
ately setose on both pairs, facial seta formula on.
article 4 = 1 and 1, on article 5 = 1 and 1; main
spine of article 5 extending to M. 75 on article 6,
article 5 lacking proximoposterior spines; spine
formula of article 6 on pereopods 1-2 = 2 + 3 plus
middistal seta, some spines long; acclivity on inner
margin of dactyls on pereopods 1-2 obsolescent,
emergent setule short, mid facial pluseta long.
Coxae 5-7 posteroventral setule formula = 1-1-1.
Articles 4-5 of pereopods 3-4 narrow, facial spine
rows sparsely developed, facial ridge formula on
article 2 of pereopods 3-5 = 0-1-1; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 53:28:25:10,
of pereopod 4 = 66:24:14:8, of pereopod 5 =
75:14:13:7, length ratios of pereopod 3 = 7 4 :
32:29:36, of pereopod 4 = 92:63:58:60, of pereo-
pod 5=120:25:25:28; article 2 of pereopod 5
reaching middle of article 4; medial apex of arti-
cle 6 coarsely combed. Posteroventral corner of
epimeron 1 rounded, posterior margin convex,
corner naked, anteroventral margin with one long
seta, posteroventral face naked; posteroventral
corner of epimeron 2 rounded-quadrate, posterior
margin weakly convex, facial ridge absent, facial
setae = 2 anteriad in tandem; posteroventral corner
of epimeron 3 rounded, posterior margin convex,
with one setule notch, ventral margin and face
naked; epimera 1-2 with setule on posterodorsal
margin set in weak notch. Urosomite 1 with lateral
setule at base of uropod 1, articulation line com-
plete, protuberant basodorsally and then forming
weak posterior saddle. Urosomite 3 weakly pro-
tuberant dorsally. Rami of uropods 1-2 with articu-
late large apical nails, outer ramus of uropod 1
with one dorsal spine, inner with one, outer ramus
of uropod 2 with one dorsal spine, inner with one;
peduncle of uropod 1 with one apicolateral spine
and 3 basofacial setae, medially with 2 ordinary
marginal spines plus thick medial spine at base of
inner ramus; peduncle of uropod 2 with 3 dorsal
spines, medially with one apical spine; apicolateral
corners of peduncles on uropods 1-2 with comb.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
XI
FIGURE 55.?Wildus mullokus, new species, holotype, male "a," S.19 mm.
NUMBER 245 141
FIGURE 56.?Wildus mullokus, new species, holotype, male "a," 3.19 mm.
142 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 57.?Wildus mullokus, new species, male "v," 3.44 mm.
Peduncle of uropod 3 with 4 ventral spines, one
elongate, dorsally with one lateral spine, one
medial spine; rami submasculine, inner extending
to M. 80 on article 1 of outer ramus, apex with
2 setae, medial margin setose, lateral margin with
one subapical seta, article 2 of outer ramus ordi-
nary, 0.22, bearing 2 medium to long setae, apico-
medial margin of article 1 with one short spine-seta,
lateral margin with 2 acclivities, spine formula =
1-1-2, setal formula = 1-1-1, medial margin setose.
Telson elongate, length-width ratio = 19:11, not
fully cleft, each apex narrow, lateral acclivity broad,
bearing ordinary lateral setule, spine next medial
longer than setule, base bearing short denticle
rows, midlateral setule pairs short. Cuticle with
ordinary sparse bulbar setules of varying sizes, sur-
face bearing fine striations in form of linear finger-
print pattern, emergent setules elongate, branched.
OBSERVATIONS.?Prebuccal parts, left mandible
and lower lip missing or damaged on holotype.
VARIATIONS.?Male "c," 4.02 mm: Outer ramus
of uropod 1 with 2 dorsal spines, peduncle with 3
NUMBER 245 143
basofacial setae, lateral apex of peduncle with one
spine. Forming intergrade between holotype and
large aberrant male "v."
Large, aberrant male "v," 3.44 mm, from type-
locality: Missing parts include: left mandible, pre-
buccal parts, right uropod 3, inner rami of uropod
1, right pereopod 3, left pereopod 1, and most of
both members of pereopod 4. This specimen prob-
ably combining features of senility and regenera-
tion; spines and setae more numerous than on
holotype except on uropod 1 basofacially, left uro-
pod 3 apparently regenerant. Head similar to that
of holotype but eye slightly larger and more elon-
gate. Article 2 of antenna 1 with 7 ventrodistal
setae; accessory flagellum 5-articulate; primary fla-
gellum 8-articulate, with 3 calceoli. Facial spine
formula on article 4 of antenna 2 = 1-6-1, dorsal
notch with 3 setal spines, dorsoproximal group
with 3 members; flagellum 25-articulate, bearing
calceolus only on article 1 or articles 1-4 (right or
left, thus aberrant or senile). Distal branch on right
lacinia mobilis broader and more distinctly bifid
than on holotype; article 2 of mandibular palp
with 3 long apical setae in tandem, article 3 with
8 apical setae. Inner plate of maxilliped lacking
medial seta; outer plate with 6 spines; article 3
of palp with one midlateral spine and 2 facial
spines. Anteroventral corner of coxa 1 bulging
more than in holotype; long setae on coxae \-A =
9-9-9-6; coxa 4 broader than in holotype. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 2-2-2-3, long anteriors = 1-2-0-0,
short anteriors = 2-2-1-1. Article 6 of gnathopod
1 larger relative to articles 4 and 5 than in holo-
type; gnathopod 2 much better developed (see illus-
tration). Pereopods 1-2 more setose than in holo-
type; article 5 with 2 and 3 apicolateral setae;
article 6 with rows of 3 and 4 spines plus midpos-
terior seta. Article 5 of pereopod 3 slightly thinner
than in holotype. Peduncle of uropod 1 with only
one basofacial seta, peduncle with 3 apicolateral
spines, 4 mediomarginal spines plus distal medial
spine at base of inner ramus; peduncle of uropod
2 with 6 dorsal spines; outer ramus of uropod 1
with 2 spines, inner ramus missing, outer ramus
of uropod 2 with 2 dorsal spines, inner naked.
Uropod 3 aberrant, probably regenerant (see illus-
tration), noting deep setal sheaths. Telson shorter
than in holotype, apicolateral setule on each lobe
elongate.
ILLUSTRATIONS.?Holotype: Top of head prob-
ably flattened slightly from preservational effects,
eyes displaced, drawn in outline only on dorsal
view.
Male "v," 3.44 mm: Views of molar and inner
plate of maxilla 1 corresponding to those of holo-
type, not duplicated for latter; minor differences
noted in palp of maxilla 1 and much broader dis-
tal branch of right lacinia mobilis; part of lower
lip reconstructed, noting short mandibular lobe;
rudimentary bulbar setule of cuticle more fre-
quently occurring than on holotype.
HOLOTYPE.?AM, male "a," 3.19 mm.
TYPE-LOCALITY.?AM P. 18125, 12 Dec 1939,
Antechamber Bay, Kangaroo Island, South Aus-
tralia, with light in net at night.
VOUCHER MATERIAL.?Type-locality, aberrant-
senile male "v," 3.44 mm (illus.); EBS 31, male
"c," 4.02 mm. Female unknown.
RELATIONSHIP.?This species differs from Wildus
thambaroo in the broader spread of the setae on
coxae 1-3, in the presence of setae on coxa 4, in
the larger distal branch of the right lacinia mobilis,
in the spine formula on the posterior margin of
article 6 on pereopods 1-2, and the highly elongate
pereopod 4. By disregarding the distal spines on
article 6 of pereopods 1-2 one counts a long seta
and a pair of stout spines, whereas in W. tham-
baroo the count is 3 long setae and 2 short spines
in tandem.
Wildus mullokus has many similarities to Brol-
gus tavelus and B. tattersalli. Wildus mullokus dif-
fers from B. tavelus in the presence of 2 apical setae
on article 2 of uropod 3, the absence of a facial
ridge on epimeron 2 and the presence of a long
seta on article 6 of pereopods 1-2. Male "v" of W.
mullokus is attached here rather than to B. tavelus
on the characters of epimeron 2 and pereopods 1-2,
although uropod 3 is aberrant and has the single
apical seta characteristic of B. tavelus.
Wildus mullokus differs from B. tattersalli in the
absence of a facial ridge on epimeron 2, the pos-
terior seta of pereopods 1-2, the presence of only
one seta on epimeron 1, and the presence of 2 api-
cal setae on article 2 of the outer ramus on uropod
3. In addition, the main setal spine on the telson
is short and thick, not elongate and seta-like.
MATERIAL.?AM, one sample (one specimen and
aberrant specimen); EBS, one sample (1).
DISTRIBUTION.?South Australia, Kangaroo Island,
144 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
neritic; New South Wales, Jervis Bay, Murray's
Beach, 6.1 m.
Wildus? fuegiensis (Schellenberg), new combination
Parharpinia fuegiensis Schellenberg, 1931:78-80. fig. 40.?
Stephensen, 1949:5-6.
Paraphoxus fuegiensis.?]. L. Barnard, 1960:271, pi. 42.
REMARKS.?This species has the definitive attri-
butes of Wildus but 3 characters might be analyzed
further as to their generic value: the presence of 3
ventral setae on epimeron 3, the odd placement of
basal setae on uropod 1 and the unusual antenna
1 of the male resembling that of male Elpeddo
kaikai. Parharpinia fuegiensis appears to differ
more strongly from Elpeddo than from Wildus in
the absence of basal spines on article 4 of antenna
2 but if the length of setae on article 2 of the outer
ramus on uropod 3 were discounted, P. fxiegiensis
could be assigned provisionally to Elpeddo. Evalua-
tion of male antenna 1 must be extended further
as this is a rarely encountered situation and may
be a recurrent aberrancy.
DISTRIBUTION.?Magellanic, 0-91 m; South Geor-
gia, 73-311 m; Tristan da Cunha, surface.
Wildus? waipiro (J. L. Barnard), new combination
Paraphoxus waipiro J. L. Barnard, 1972b: 143-145, figs. 79,
80.
REMARKS.?This species is distinguished in the
key to Wildus, but several characters must be eval-
uated with fresh material; the presence of a special
apicomedial spine on the peduncle of uropod 1
is not confirmed.
DISTRIBUTION.?New Zealand, intertidal.
Paraphoxus Sars
Paraphoxus Sars, 1895:148-149.?Stebbing, 1906:137.?Bar-
nard and Drummond, 1976:524.
Not Pontharpinia Stebbing, Parharpinia Stebbing, Proto-
phoxus K. H. Barnard, Trichophoxus K. H. Barnard, nor
Metharpinia Schellenberg.
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae confined apically.
Article 1 of antenna 2 not ensiform, article 3 with
2 setules, facial spines on article 4 in one main row,
with rudimentary second row, article 5 short. Right
mandibular incisor with 3 teeth; molar not tritura-
tive, small, conical, bearing 3 long clumped spines
with common base, not bearing fuzz; palpar hump
small. Palp of maxilla 1 biarticulate, inner plate
with 2 setae. Setation of maxilla 2 ordinary. Inner
plate of maxillipeds ordinary; apex of palp article
3 weakly protuberant, dactyl elongate, apical nail
distinct, elongate. Gnathopods small, similar, gnath-
opod 2 weakly enlarged; article 5 of gnathopods
1-2 very short, almost cryptic, with eusirid attach-
ment; palms oblique, hands of gnathopods 1-2
weakly enlarged, ovatorectangular, elongate, poorly
setose anteriorly. Article 5 of pereopods 1-2 setose
posteroproximally. Article 2 of pereopod 3 of broad
form, articles 4-5 of pereopods 3-4 narrow, article
2 of pereopods 3-4 not setose posteriorly; pereopod
5 ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Epimera 1-2 lacking long pos-
terior setae, without midfacial setae above ventral
facial ridge; epimeron 3 of rounded classification
and lacking long setae. Urosomite I generally
naked; urosomite 3 without dorsal hook or special
process. Peduncle of uropod 1 normally elongate,
without apicoventral spike, without special en-
larged apicolateral-medial spine; peduncular apices
of uropods 1-2 combed; spine(s) on inner ramus
of uropod 1 in one row, no rami continuously spi-
nose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 especially elongate, article 2 of outer
ramus carrying 2 short to medium apical setae.
Telson ordinary, with only one apical spine on
each lobe plus setules.
DESCRIPTION.?Rostrum fully developed. Fuzz on
article 1 of antenna 1 in male present; calceoli on
male primary flagellum of antenna 1 present. Cal-
ceoli on article 5 of male antenna 2 present; flagel-
lum in male with calceoli. Prebuccal parts ordi-
nary, extended forward, massive, poorly separated
from each other, upper lip dominant. Right lacinia
mobilis bifid, flabellate; mandibular palp thin, arti-
cle 1 slightly elongate, article 2 without outer setae,
apex of article 3 almost transverse. Lower lip bear-
ing cones. Outer plate of maxilla 1 with 11 spines,
one spine especially thickened. Inner plates of max-
illiped poorly armed, thick. Coxae 2-4 without spe-
cial anterodorsal humps. All posterior spines on
article 6 of pereopods 1-2 thin and stiff, midapical
spine or seta absent; dactyls of pereopods 1-2 lack-
ing inner tooth, bearing raphe, apical nail distinct
NUMBER 245 145
but immersed. Article 2 of pereopod 5 without
facial setae. Peduncle of uropod 1 with dorsolateral
spines confined apically, medial spines widely
spread; peduncle of uropod 2 with medial spines
and setae sparse but widely spread. Peduncle of
uropod 3 lacking extra subapical setae or spines.
Telson with ordinary pair of midlateral or dorsal
setules on each side.
TYPE-SPECIES.?Phoxus oculatus Sars, 1879 (mon-
otypy).
COMPOSITION.?Possibly Parharpinia simplex Gur-
janova, 1938.
RELATIONSHIP.?Paraphoxus differs from Brolgus
and Ganba in the presence of two apical setae on
the outer ramus of uropod 3. It differs from Kuri-
tus, Elpeddo and Wildus in the absence of the spe-
cial apicomedial spine on the peduncle of uropod
1.
See Barnard and Drummond (1976) for an expo-
sition on the nomenclatural history of Paraphoxus,
the revival of its synonyms, and notes on the place-
ment of species to be rejected from Paraphoxus.
Also see "Allocation of Species in Paraphoxus"
below.
Paraphoxus oculatus (Sars)
Phoxus oculatus Sars, 1879:441; 1885:154-156, pi. 13: figs.
4. 4a-e.
Paraphoxus oculatus.?Stebbing, 1906:137-138 [with refer-
ences]. -Oldevig, 1959:S8.-J. L. Barnard, 1960:240-243,
pis. 27, 28 [with references]; 1966a:89; 1967:138; 1971:
7O.-Nagata, 1965:156.?Karaman, 1973:91-98, figs. 25-
28.?Griffiths, 1974b:321.
Phoxus maculatus Chevreux, 1888:2-3.
Paraphoxus maculatus.?Chexreux, 1900:34-36, pi. 5: fig. 5;
1911:187, pi. 10: figs. 12, 13.-Chevreux and Fage, 1925:
103-104, fig. 97.?Ledoyer, 1968:191.
DISTRIBUTION.?Atlantic and North Pacific
Oceans, to depths of 860 m.
Paraphoxus? simplex (Gurjanova)
Parharpinia simplex Gurjanova, 1938:272-274, 385-386, fig.
10.
Pararpinia [sic] simplex.?Gurjanova, 1951:392-394, fig. 238.
Paraphoxus simplex.?J. L. Barnard, 1960:278.
REMARKS.?Many characters of this species need
confirmation for its relegation to Paraphoxus; one
possible mark of its distinction from Paraphoxus
oculatus is the stoutness of spines on article 6 of
pereopods 1-2.
DISTRIBUTION.?Japan Sea.
Allocation of Species in Paraphoxus
Paraphoxus abronius J. L. Barnard, 1960, to a genus to be
described.
Protophoxus australis K. H. Barnard, 1930, to Protophoxus.
Pontharpinia barnardi Pirlot, 1932, to a genus to be de-
scribed.
Phoxus batei Haswell, 1879, to Birubius.
Paraphoxus bicuspidatus J. L. Barnard, 1960, to a genus to
be described.
Pontharpinia calcarata Gurjanova, 1938, to a genus to be de-
scribed.
Trichophoxtis capillatus K. H. Barnard, 1930, to Tricho-
phoxus.
Pontharpinia centralis Schellenberg, 1938, to a genus to be
described.
Paraphoxus chelatus Cooper, 1974, to a genus to be de-
scribed.
Paraphoxus cognatus J. L. Barnard, 1960, to a genus to be
described.
Metharpinia cornuta Schellenberg, 1931, to Metharpinia.
Paraphoxus daboius J. L. Barnard, 1960 to a genus to be
described.
Pontharpinia epistoma Shoemaker, 1938, to a genus to be
described.
Paraphoxus fatigans J. L. Barnard, 1960, to a genus to be
described.
Pontharpinia floridana Shoemaker, 1933, to a genus to be
described.
Parharpinia fuegiensis Schellenberg, 1931, to Wildus.
Paraphoxus gemmatus J. L. Barnard, 1964a, to a genus to be
described.
Phoxus geniculatus Stimpson, 1856, dubious.
Phoxus grandis Stimpson, 1857, dubious.
Paraphoxus heterocuspidatus J. L. Barnard, 1960, to a genus
to be described.
Pontharpinia intermedia Schellenberg, 1925, to Basuto stimp-
soni.
Paraphoxus jonesi J. L. Barnard, 1963, to a genus to be de-
scribed.
Metharpinia longirostris Schellenberg, 1931, to Metharpinia.
Paraphoxus lucubrans J. L. Barnard, 1960, to a genus to be
described.
Phoxus maculatus Chevreux, 1888, to Paraphoxus oculatus.
Pontharpinia maxima Stephensen, 1947, possibly to Para-
phoxus pyripes.
Pontharpinia milleri Thorsteinson, 1941, to a genus to be
described.
Pontarpinia [sic] nasuta Gurjanova, 1936, to a genus to be
described.
Parharpinia obliqua K. H. Barnard, 1932, to a genus to be
described.
Pontharpinia obtusidens Alderman, 1936, to a genus to be
described.
146 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Phoxus obtusus Stimpson, 1855, dubious.
Pararpinia [sic] ochotica Gurjanova, 1953, to a genus to be
described.
Urothoe pinguis Haswell, 1880, to Pontharpinia.
Parophoxus pyripes K. H. Barnard, 1930, to a genus to
described.
Paraphoxus rakiura Cooper and Fincham, 1974, to Waitangi.
Paraphoxus robustus Holmes, 1908, to a genus to be de-
scribed.
Urothoe rostratus Dana, 1853, to a genus to be described or
Birubius.
Parharpinia rotundifrons K. H. Barnard, 1932, to a genus to
be described.
Paraphoxus similis J. L. Barnard, I960, to a genus to be
described.
Parharpinia sinuata K. H. Barnard, 1932, to a genus to be
described.
Paraphoxus spinibasus Cooper, 1974, to a genus to be de-
scribed.
Paraphoxus spinosus Holmes, 1908, to a genus to be de-
scribed.
Paraphoxus stebbingi J. L. Barnard, 1958. to Tipimegus.
Paraphoxus stenodes J. L. Barnard, 1960, to a genus to be
described.
Pontharpinia stimpsoni Stebbing, 1908, to Basuto.
Paraphoxus subuncigerus Kudrjaschov, 1965, to a genus to
be described.
Paraphoxus tattersalli J. L. Barnard, 1958, to Brolgus.
Pontharpinia tridentata J. L. Barnard, 1954, to a genus to be
described.
Parharpinia uncigera Gurjanova, 1938, to a genus to be de-
scribed.
Pontharpinia uncinata Chevreux, 1912a,b, to a genus to be
described.
Phoxus unicirostratus Giles, 1890, to Mandibulophoxus.
Paraphoxus variatus J. L. Barnard, 1960, to a genus to be
described.
Paraphoxus vigitegus J. L. Barnard, 1971, to a genus to be
described.
Phoxus villosus Haswell, 1879, to Parharpinia.
Paraphoxus waipiro J. L. Barnard, 1972b, to Wildus.
Paraphoxus sp. Australia, J. L. Barnard, 1960, to Matong
matong.
Paraphoxus sp. South Africa, J. L. Barnard, 1960, dubious.
Pontharpinia sp., Falkland Islands, (= P. rostratus, Stebbing,
1914), dubious.
Pontharpinia sp., Flores Sea, (= P. rostrata, Pirlot, 1932, fig.
15 only), dubious.
LEONGATHINAE
DIAGNOSIS.?Article 2 of antenna 1 ordinary;
mandibular molar semitriturative, bearing one ex-
tended and toothed margin; palp of maxilla 1
biarticulate; setation on maxilla 2 ordinary; gnatho-
pod 2 significantly enlarged; article 2 of pereopod
3 of broad form, tapering distally; pereopod 5
ordinary.
DESCRIPTION.?Article 5 of antenna 2 reduced
in size; epimeron 3 of nonrounded classification;
apices of peduncles on uropods 1-2 not combed.
TYPE GENUS.?I.eongathns, new genus.
COMPOSITION.?Unique.
Leongathus, new genus
DIAGNOSIS.?Eyes obsolescent. Flagella of anten-
nae 1-2 unreduced in female. Article 2 of antenna
1 ordinary, ventral setae confined proximally.
Article 1 of antenna 2 weakly ensiform; article 3
with 2 setules; facial spines on article 4 in 2 in-
distinct rows, plus special apical spines; article 5
short. Right mandibular incisor with S teeth;
molar partly triturative, medium broad, conical,
bearing 7 or more splayed nonarticulate teeth,
bearing fu/z; palfMi hump medium. Palp of maxilla
1 biarticulate; inner plate with I setae. Setation
of maxilla 2 ordinary. Inner plate of maxillipeds
ordinary; apex of palp article 3 not or weakly pro-
tuberant, dactyl elongate, apical nail mostly im-
mersed, short. Gnathopods highly dissimilar,
gnathopod 2 strongly enlarged; article 5 of gnatho-
pod 2 very short, cryptic, ordinary on gnathopod 1,
without eusirid attachment; palms oblique, hand
of gnathopods 1-2 elongate and broadened respec-
tively, poorly setose anteriorly. Article 5 of pereo-
pods 1-2 with posteroproximal setae. Article 2 of
pereopod 3 of broad form but tapering distally,
articles 4-5 of pereopods 3-4 narrow to medium,
article 2 of pereopod 4 setose posteriorly; pereopod
5 ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Epimera 1-2 bearing numerous
posterior setae, without midfacial setae above ven-
tral facial ridge; epimeron 3 ordinary. Urosomite 1
bearing one or more midvemral crescents or bun-
dles of setae; urosomite 3 without dorsal hook or
special process. Peduncle of uropod 1 normally elon-
gate, without apicoventral spike, without special en-
larged apicolateral-medial spine; peduncular apices
of uropods 1-2 not combed; uropod 2 with long setae
on peduncle, spines if present on inner ramus of uro-
pod 1 in one row, no rami continuously spinose to
apex, inner ramus of uropod 2 ordinary. Uropod 3
ordinary, article 2 of outer ramus carrying 2 long
apical setae. Telson ordinary, with only one apical
spine or seta on each lobe plus setules.
DESCRIPTION.?Rostrum fully developed, uncon-
stricted. [Fuzz on article 1 of antenna 1 in male
NUMBER 245 147
unknown, calceoli on male antenna 1 and antenna
2 unknown.] Ventral setules on article 1 of antenna
1 widely spread. Prebuccal parts ordinary, poorly
separated from each other, upper lip dominant.
Right lacinia mobilis bifid, distal branch flabellate;
mandibular palp medium to thin, article 1 short,
article 2 with outer setae, apex of article 3 oblique.
Lower lip bearing cones. Outer plate of maxilla 1
with 11 spines, one spine especially thickened.
Inner plates of maxilliped thick, ordinarily setose.
Only coxa 2 with special anterodorsal hump. Some
posterior spines on article 6 of pereopods 1-2 thin
and seta-like, midapical spine or seta absent. Article
2 of pereopod 5 with facial setae. Peduncle of uro-
pod 1 with dorsolateral spines confined apically,
basofacial setae dispersed, medial spines widely
spread; peduncle of uropod 2 with only one medial
spine confined apically. Peduncle of uropod 3 lack-
ing extra subapical setae or spines. Telson with
ordinary pair of midlateral or dorsal setules on
each side.
TYPE-SPECIES.?Leongathus nootoo, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?The mandibular molar of
Leongathus appears to be transitional between the
fully triturative form of Pontharpinia and the
spinose kind of the Matong and Birubius groups of
genera.
Leongathus differs from Pontharpinia in the
apomorphic right lacinia mobilis bearing a fully
fused proximal branch and in the ordinary kinds
of maxilla 1, telson, article 3 of antenna 2, labial
cones, dactyl of pereopod 5, article 5 of pereopods
1-2, reduced rows of spines on article 4 of antenna
2, tapering article 2 of pereopod 3, fewer facial
setae on article 2 of pereopod 5, these not formed
into bundles, apically confined dorsal spines on the
peduncle of uropod 1, widely dispersed basofacial
setae on uropod 1, and absence of the special pe-
duncular spine, presence of posterior setae on epi-
mera 1-2, and elongation of peduncular spines on
uropod 2.
From Matong, Kotla, and Yammacoona, Leon-
gathus differs in the unusual molar, the poor nail
of the maxillipedal palp, the much shortened, al-
most vestigial article 5 of gnathopod 2, the taper-
ing article 2 of pereopod 3, the facial setae on
article 2 of pereopod 5, the normal coxae 1-3, the
elongate setal-spines on the peduncle of uropod 2,
and the numerous and dispersed basofacial setae
on uropod 1. Leongathus furthermore differs from
Matong and Kotla in the thick antenna 1 with
proximal ventral setae on article 2, the long and
free inner ramus of uropod 2, and in the confine-
ment of most spines on article 4 of antenna 2 into
one row. Leongathus also differs from Yammacoona
in the normal plates of maxilliped, absence of a
facial brush on epimeron 3, presence of posterior
setae on epimera 1-2 and the wide spread of me-
dial spines on the peduncle of uropod 1.
Leongathus differs from Mandibulophoxus and
Basuto in the smaller palpar hump on the mandi-
ble, the well developed molar, the absence of a
special peduncular spine on uropod 1, the more
elongate article 2 of antenna 1 with the ventral
setae situated proximally, the weak nail on the
maxilliped, and the noncontinuous dorsal spina-
tion on the rami of uropods 1-2.
Leongathus nootoo, new species
FICURES 58-60
DESCRIPTION OF FEMALE.?Head about 17 percent
of total body length, greatest width about 55-60
percent of length; rostrum unconstricted, elongate,
almost reaching apex of article 2 on antenna 1.
Eyes small, apparently present, clear of pigment,
ommatidia apparently absent. Article 1 on pe-
duncle of antenna 1 about 1.5 times as long as
wide, about twice as wide as article 2, ventral mar-
gin with about 7 setules, weakly produced dorsal
apex with 2 setules; article 2 about 0.5 times as
long as article 1, with ventral, proximal cycle of
3-6 setae, primary flagellum with 9-10 articles,
about 0.45 times as long as peduncle, bearing short
aesthetascs; accessory flagellum with 7-8 articles.
Spine formula on article 4 of antenna 2 = 1-4-4
or 1-3-6 or 1-3-5, dorsal margin with one proxi-
mal seta and notch bearing 2 setae and one spine,
ventral margin densely setose distally, with 2 ven-
trodistal long to medium spines; article 5 about 0.7
times as long as article 4, facial spine formula =
1-1 or 1-1-1, dorsal margin naked, ventral margin
densely setose, with 2 ventrodistal long to medium
spines, one of these subdistal and facial; flagellum
about 0.8 times as long as articles 4-5 of peduncle
combined, with 8-9 articles. Mandibles with me-
dium palpar hump; right incisor with 3 weak teeth;
left incisor with 3 humps in 2 branches; right la-
148 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 58.?Leongathus nootoo, new species, holotypc, female "a," 7.4 mm (w <m female "w,1
8.6 mm).
NUMBER 245 149
FIGURE 59.?Leongathus nootoo, new species, holotype, female "a," 7.4 mm.
cinia mobilis bifid, distal branch shorter than
proximal, flabellate, denticulate, proximal branch
simple, pointed, with marginal denticles; left la-
cinia mobilis with 5 adz-teeth plus accessory den-
ticulations; right rakers 7-9 plus 1-2 rudimenta-
ries; left rakers 8-9 plus 2 + rudimentaries; molars
composed of elongate lamina in form of soft cone
bearing 6-7 plus rudimentary cusps and one short
plusetule weakly disjunct, molars fuzzy; palp article
1 short, article 2 with 3 long to medium inner api-
cal setae, 5 other long and short inner setae and 3
outer facial and one outer marginal setae, article 3
about 1.1 times as long as article 2, oblique apex
with 9 spine-setae, basofacial formula = (l-l)-l .
Inner plate of maxilla 1 ordinary, bearing one long
apical pluseta, one longer apicomedial seta, 2 apico-
lateral much shorter setae; palp article 2 with 4-5
apical-medial marginal spines and 5 apicolateral
to medial setae. Plates of maxilla 2 extending sub-
equally, outer much broader than inner, outer with
3 apicolateral setae, inner with one medial seta.
Inner plate of maxilliped with one large apical
spine, 3 apicofacial and one ventrodistal setae, 2
medial setae; outer plate very large, with 10 medial
and apical spines, 6 apicolateral setae; palp article
1 lacking apicolateral setae, article 2 with one api-
colateral seta, medial margin of article 2 weakly
setose, article 3 unprotuberant, with one facial seta,
2 lateral setae, nail of article 4 obsolescent, mostly
immersed, short, with 2 accessory setules. Coxa 1
scarcely expanded distally, anterior margin weakly
concave; main ventral setae of coxae 1-4 = (15?18)?
(11?15)?(11?14)?18, additional posteriormost setule
of coxae 1-3 shortest; anterior and posterior mar-
gins of coxa 4 parallel, posterior margin convex to
straight, posterodorsal corner rounded, posterodor-
sal margin short, V-shaped, width-length ratio of
coxa 4 = 16 : 17. Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 = (1-2)-
(3-5)-2-3, long anteriors = (5-6)-8-(4-5)-3, short
posteriors = 1-1-0-0, no others. Gnathopod 1 ordi-
nary, gnathopod 2 enlarged; article 5 tiny, cryptic-
article 6 very broad and elongate; width ratios of
articles 5-6 on gnathopods 1-2 = 30:32 and 28:60,
150 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 60.?Leongathus nootoo, new species, holotype, female "a," 7.4 mm (w -? female "w,"
8.6 mm).
NUMBER 245 151
length ratios = 56:72 and 14:110; palmar hump
of gnathopod 2 very large, palms strongly oblique;
article 5 of gnathopod 1 ovate, posterior margin
rounded; article 5 of gnathopod 2 short, triangular,
lobate. Pereopods 1-2 similar; facial seta formula
on article 4 ? 3-6 and 3-6, on article 5 = 5-7 and
5-7; main spine of article 5 extending to M. 100
on article 6, article 5 lacking proximoposterior
spines; spine formula of article 6 = 4 + 4 or 5 +5 ,
no middistal seta, most spines especially long and
thin; acclivity on inner margin of dactyls of pe-
reopods 1-2 weak, obsolescent, emergent setule
short, almost fully immersed, midfacial pluseta ordi-
nary. Coxae 5-7 posteroventral seta formula =
(8-11)-(2-3)-(5-5). Articles 4-5 of pereopods 3-4
narrow, facial spine rows sparse, facial ridge for-
mula on article 2 of pereopods 3-5 = 0-1-2, ante-
rior ridge on pereopod 5 short, article 2 of pereo-
pod 3 tapering distally; width ratios of articles 2, 4,
5, 6 of pereopod 3 =46:23:22:14, of pereopod
4 = 65:30:24:14, of pereopod 5 = 88:23:16:9, length
ratios of pereopod 3 = 64:26:34:38, of pereopod
4 = 82:55:58:62, of pereopod 5 = 100:30:22:30; arti-
cle 2 pf pereopod 5 almost reaching apex of article
4, with row of posterofacial setae; medial apex of
article 6 with finely combed hump. Posteroventral
corner of epimeron 1 rounded, posterior margin
straight to weakly convex, setose, postero- and
anteroventral margins continuously setose; postero-
ventral corner of epimeron 2 with small, sharp
tooth guarded by sinus, posterior margin weakly
convex, serrate, setose, facial setae = dense and
scattered, some pairs set vertically; posteroventral
corner of epimeron 3 with medium to large tooth,
posterior margin almost straight, weakly serrate,
setose, ventral margin with 4-5 facial setae mainly
in middle. Urosomite 1 with or without lateral
setule at base of uropod 1 plus 3-5 ventral setae
in group, with or without additional 1-2 setae
more anteriorly, articulation line complete; uro-
somite 3 weakly protuberant dorsally. Rami of
uropods 1-2 with articulate apical nails, outer
ramus of uropod 1 with 1-2 dorsal spines, inner
with one, outer ramus of uropod 2 with 2 dorsal
spines, inner lacking dorsomedial spines; peduncle
of uropod 1 with one apicolateral spine and sets
of 1-3-3 or 1-2 basofacial setae, medially with 8
marginal setae and spines; peduncle of uropod
2 with 7-10 long dorsal setae and one apical spine,
medially with one small apical spine; apicolateral
corners of peduncles on uropods 1-2 lacking comb.
Peduncle of uropod 3 with 4-5 ventral spines,
dorsally with one lateral spine and one setule,
one medial spine and 2 setules; rami feminine,
inner extending to M. 60 on article 1 of outer
ramus, apex with one short seta, medial and lateral
margins naked, article 2 of outer ramus short, 0.14,
bearing 2 long to medium setae, apicomedial mar-
gin of article 1 with 3 spine-setae, lateral margin
with 1-2 acclivities, spine formula = 1-3-4 or
0-2-3, setal formula = 0. Telson short, length-
width ratio = 2:3, not fully cleft, each apex wide,
rounded, lateral acclivity narrow, weak, bearing
short lateral setule, spine-seta next medial much
longer than setule, midlateral setules diverse. Cuti-
cle with ordinary bulbar setules mixed with pipes,
emergent setules especially branched.
OBSERVATIONS.?Eyes composed of weakly visible
capsule of irregular tissue (marked in dashed out-
line in lateral view of head), not visible in dorsal
view owing to thickness of cuticle. Description
based on both adult females in hand but following
parts of female V not examined: maxillae 1-2,
maxilliped, upper and lower lips, epistome.
Juvenile, 4.1 mm: Generally like adult but
armaments fewer, for example: article 2 of antenna
1 with 2 ventral setae, primary flagellum with 6
articles, accessory flagellum with 5 articles; spine
formula on article 4 of antenna 2 = 1-2-1, article
5 with one facial spine, flagellum 6-articulate; long
setae on coxae 1, 2, 4, = 6, 5, 4; epimeron 1 with
only 3 setae, one of these anteroventral, one pos-
teroventral, one ventral; epimeron 2 with 3 lateral
setae; epimeron 3 with 4-5 posterior setae and one
ventral seta; urosomite 1 with one seta at base of
uropod 1, ventral setae present; peduncle of uro-
pod 2 with 3 setae and one spine; basofacial setae
of uropod 1 = 1-1; outer ramus of uropod 3 with
0-1 lateral spine.
ILLUSTRATIONS.?One seta on article 2 of antenna
1 missing (6 instead of 5); lower lip drawn from
oral side, inner lobes cleft on aboral side; overall
view of right mandible showing unflattened ver-
sion; on overall view of left mandible those raker
spines situated behind molar omitted; enlarged
views of right mandibular molars from 2 different
specimens drawn from opposite sides, thus appear-
ing as mirror images; several spines missing on
posteroventral corner of article 4 on pereopod 4;
epimera of holotype not illustrated, epimeron 1
152 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
with 18 setae, epimeron 2 with 11 setae, epimeron
3 with 12 posterior and 4 ventral setae;
HOLOTYPE.?AM, female "a," 7.4 mm.
TYPE-LOCALITY.?AM P. 18215, 18 Jun 1962, off
Sydney, New South Wales, Australia, 33?58.4' S,
151 ?29' E, 150 m.
VOUCHER MATERIAL.?Type-locality: female "w,"
8.6 mm (illus.); AM P. 18248, juvenile "c," 4.1 mm
(only other specimen known). Male unknown.
MATERIAL.?AM, 2 samples (3).
DISTRIBUTION.?New South Wales, Sydney, 150 m.
JOUBINELLINAE
DIAGNOSIS.?Article 2 of antenna 1 ordinary to
elongate; mandibular molar fully triturative or re-
duced to a small hump with articulate spines; palp
of maxilla 1 biarticulate; setation on maxilla 2
ordinary to reduced; gnathopod 2 significantly en-
larged; article 2 of pereopod 3 of broad form, not
tapering distally; pereopod 5 ordinary.
DESCRIPTION.?Article 5 of antenna 2 of normal
size or rarely reduced; epimeron 3 of nonrounded
classification; apices of peduncles on uropods 1-2
not combed.
TYPE GENUS.?Joubinclla Chevreux.
COMPOSITION.?Kotla, new genus; Matong, new
genus; Yammacoona, new genus, and the "univer-
sal" genus Cunmuura, new genus, placed provision-
ally in the Brolginae.
REMARKS.?The members of this subfamily are
highly diverse. Joubinella is a pelagic genus bearing
eusirid gnathopods and possible predatorial adapta-
tions, whereas Kotla, Matong, and Yammacoona
appear to have benthic inquilinous adaptations in
reduced rostra, thickened and oddly spinose anten-
nae, reduction in uropod 3, fusion to the peduncle
of the inner ramus of uropod 2 and in some cases
nonskid cuticle. Cunmurra is a transitional genus,
having affinities mainly to the Brolginae but form-
ing an intergradc towards the Birubius mayamayi
group of species and being convergent towards the
Phoxocephalinae. Because of the tendency to
identify Cunmurra with the brolgtns in its overall
appearance we have removed it to a position in
that subfamily. In its gnathopods, head, epimera
and antennae, Cunmurra appears closer to the
hypothetical primitive joubinellin than any of the
other genera so contained.
Key to the Genera of Joubinellinae
1. Gnathopods of highly eusirid, pelagic, and predatorial form, gnathopod 1 usually much
larger than gnathopod 2 but occasionally of equal size, antenna 2 extremely thin, lacking
well organized clusters of facial spines, flagellum in female reduced to 2-3 articles, mandi-
bular molar strongly triturative Joubinella
Gnathopods weakly eusirid, of nonpelagic and nonpredatorial form, gnathopod 2 usually
much larger than gnathopod 1 but occasionally of equal size, antenna 2 stout to medium
in thickness, bearing well organized clusters of facial spines, flagellum in female exceeding
6 articles, mandibular molar not triturative 2
2. Epimeron 3 reduced to fully rounded classification, lacking all but 2 fully developed setae
[head normal, uropod 2 freely articulate, uropods 1-2 lacking accessory apical nails, no
epimera with facial brushes of setae above lateral ridges, spines on uropod 2 normal,
ventral setae on article 2 of antenna 1 set in middle]
Cunmurra, new genus (see Brolginae)
Epimeron 3 with 5 or more fully developed setae 3
3. Article 4 of antenna 2 bearing only 2 rows of facial and apical spines, head with rostrum
obsolete but broadly truncate from lateral view, inner ramus of uropod 2 freely articulate,
ventral setae on article 2 of antenna 1 set proximally Yammacoona, new genus
Article 4 of antenna 2 bearing 3 or more rows of facial and apical spines, head with very
short rostrum but anterior margin from lateral view sinuous, inner ramus of uropod 2
partially to fully fused to peduncle, ventral setae on article 2 of antenna I set distally.. 4
4. Spines on uropod 2 conical, of normal dimensions, sharp, epimera 1-2 lacking large vertically
set setal brushes Matong, new genus
Spines on uropod 2 of rounded, jewel-like form, blunt, epimera 1-2 bearing large vertically
set setal brushes Kotla, new genus
NUMBER 245 153
Joubinella Chevreux
Joubindla Chevreux, 1908:8-11.?J. L. Barnard, 1960:357-
358.
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 reduced in female. Article 2 of antenna 1
elongate to ordinary, ventral setae absent or con-
fined apically. Article 1 of antenna 2 not ensiform,
article 3 with 2 setules; facial spines on article 4
variable, absent (type) or in 2 or more rows; article
5 ordinary in size to especially thin. [PRight mandi-
bular incisor with 2-4 teeth]; molar strongly tritur-
ative, medium; palpar hump small. Palp of maxilla
1 biarticulate; inner plate naked. Setation of
maxilla 2 ordinary (type) to weak. Inner plate of
maxillipeds ordinary; apex of palp article 3 not
protuberant, dactyl elongate, apical nail distinct,
stout, medium. Gnathopods dissimilar, moderately
to strongly enlarged; article 5 of gnathopod 2 of
ordinary length, free or almost cryptic, highly elon-
gate on gnathopod I, with eusirid attachment;
palms transverse to chelate, hand of gnathopods
1-2 broadened, poorly setose anteriorly. Article 5
of pereopods 1-2 setose posteroproximally. Article
2 of pereopod 3 of broad form but tapering distally;
articles 4-5 of pereopods 3-4 narrow to medium,
article 2 of pereopods 3-4 not setose posteriorly;
pereopod 5 ordinary, article 2 naked ventrally,
articles 3-4 especially small, dactyl normal. Epimera
1-2 lacking long posterior setae, without midfacial
setae above ventral facial ridge; epimeron 3 of
rounded classification and lacking long setae. Uro-
somite 1 generally naked; urosomite 3 without
dorsal hook or special process. Peduncle of uropod
1 normally elongate to slightly shortened (type),
with or without apicoventral spike, without special
enlarged apicolateral-medial spine; [Ppeduncular
apices of uropods 1-2 not combed]; inner ramus of
uropod 1 with marginal spines in one row, either
none or some rami continuously spinose to apex,
inner ramus of uropod 2 ordinary. Uropod 3 ordin-
ary, article 2 of outer ramus carrying 0-2 long to
vestigial apical setae. Telson ordinary, with 2?4
apical spines or setae on each lobe plus setules.
DESCRIPTION.?Rostrum fully developed. [Fuzz on
article 1 of antenna 1 in male unknown, calceoli on
male antennae 1-2 unknown. Prebuccal parts un-
known. PRight lacinia mobilis bifid]; mandibular
palp thin, article 1 short to slightly elongate, article
2 without outer setae, apex of article 3 oblique.
[Lower lip unknown.] Outer plate of maxilla 1 with
9 spines, one or no spine especially thickened. Inner
and outer plates of maxilliped poorly armed, thick.
Coxae 2-A without special anterodorsal humps.
Some posterior spines on article 6 of pereopods 1-2
thin and seta-like, others thick and stiff, [midapical
spine or seta unidentified]. Article 2 of pereopod 5
without facial setae. Peduncle of uropod 1 with
dorsolateral spines widely spread, medial spines
widely spread; [Ppeduncle of uropod 2 with only
one medial spine or setule confined apically or
spines widely spread]. Peduncle of uropod 3 bear-
ing or lacking extra subapical setae or spines. Tel-
son with pair of midlateral or dorsal setules on each
side, highly apical.
TYPE-SPECIES.?Joubinella ciliata Chevreux, 1908
(monotypy).
COMPOSITION.?Joubinella bychovskii Gurjanova,
1952; / . strelkovi Gurjanova, 1952; / . traditor Pirlot,
1932; / . tzvetkovae Kudrjaschov, 1965 (Okhotsk Sea,
88 m). See J. L. Barnard (1960) for other literature
and distributions.
REMARKS.?The diversity in antennae and uro-
pods suggests that at least two genera are mixed
into this complex. More elaboration of detail is re-
quired in several of the species. The diagnosis and
description for the genus are drawn from the sev-
eral species and may be confounded by this com-
bining of facts not limited to the type-species.
Key to the Species of Joubinella
1. Inner ramus of uropod 1 bearing only one dorsal spine, outer ramus of uropod 2 lacking
accessory- apical spine 2
Inner ramus of uropod 1 with 3 or more dorsal spines, outer ramus of uropod 2 with acces-
sory apical spine 3
2. Article 4 of female antenna 2 with naked dorsodistal tooth, inner ramus of female uropod 3
more than half as long as article 1 of outer ramus / . bychovskii
Article 4 of female antenna 2 with articulate spine but no tooth, inner ramus of female
uropod 3 less than half as long as article 1 of outer ramus / . tzvetkovae
154 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
S. Apicolateral spine on peduncle of uropori 1 less than one fourth as long as outer ramus,
each apex of telson with 2-3 very long spine-setae and one stout short spine, lateral surface
area of article 6 on gnathopod 1 subequal to area on gnathopod 2, article 2 of antenna
1 half as long as article 1 /? *trelkovi
Apicolateral spine on peduncle of uropod 1 half as long as outer ramus, each apex of telson
with 2-3 very short thin spines or setae, lateral surface area of article 6 on gnathopod 1
much larger than area on gnathopod 2, article 2 of antenna 1 more than two thirds as
long as article 1 *
4. Apices of telson blunt, strong setosity present on anterodistal margins of article 4 on pere-
opods 4-5 /? ciliata
Apices of telson sharp, weak setosity present on anterodistal margins of article 4 on pere-
opods 4-5 1- tradiior
Joubinella species 1
MATERIAL.?AM P. 18219, off Sydney, 33?58.4' S,
115?29' E, 150 m, juvenile, 2.25 mm.
REMARKS.?A damaged specimen not worthy of
specific description but very similar in many re-
spects to the ordinary joubinellas heretofore de-
scribed in the literature. Article 2 of uropod 3 elon-
gate, bearing 2 short apical setae; uropods 1-2 with
long apical spine-nails on rami, with one accessory
nail and with subdistal dorsal spine, the degree of
apicality indicating youth of the specimen; article 2
of antenna 1 scarcely overriding article 3; article 6
of pereopods 1-2 with spine rows of 1 + 1, spines
longer than dactyl; gnathopod 1 definitely much
larger than gnathopod 2 and with unusual shape
typical of Joubinella.
Joubinella species 2
MATERIAL.?SBS 3 (one shriveled specimen).
REMARKS.?This specimen, perhaps in the range
of 4-6 mm in length, is clearly a member of Joubi-
nella but too poorly preserved for analysis.
Matong, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae 1-2
unreduced in female. Article 2 of antenna 1 elon-
gate, ventral setae mainly apical. Article 1 of an-
tenna 2 weakly ensiform; article 3 with 2 setules;
facial spines on article 4 in 2 or more rows; article
5 ordinary in size. Right mandibular incisor with 3
teeth; molar not triturative, ordinary, medium, pil-
low-shaped, bearing 4 or more splayed, semiarticu-
late spines, usually bearing fuzz; palpar hump
small. Palp of maxilla 1 biarticulate; inner plate
with 3-4 setae. Setation of maxilla 2 ordinary. In-
ner plate of maxillipeds ordinary; apex of palp
article 3 not protuberant, dactyl elongate, apical
nail distinct, medium. Cnathopods enlarged, gna-
thopod 2 strongly enlarged; article 5 of gnathopods
1-2 of ordinary length but almost cryptic on gna-
thopod 2, without eusirid attachment; palms trans-
verse, hands of gnathopods 1-2 rectangular, broad-
ened, poorly setose anteriorly. Article 2 of pereopod
3 of broad form, articles 4-5 of pereopods 3-4
broad to medium, article 2 of pereopods 3-4 not
setose posteriorly; pereopod 5 ordinary, article 2
naked ventrally, article 3 ordinary, dactyl normal.
Epimera 1-2 bearing few short posterior setae, with-
out midfacial setae above ventral facial ridge; epi-
meron 3 ordinary. Urosomite 1 with small lateral
facial spines, lacking midventral bundles of setae,
with sparse apicoventral setae or spines near base
of uropod 1; urosomite 3 without dorsal hook or
special process. Peduncle of uropod 1 normally
elongate, without apicoventral spike, without spe-
cial enlarged apicolateral-medial spine; peduncular
apices of uropods 1-2 not combed; inner ramus of
uropod 1 with marginal spines in one row, no rami
continuously spinose to apex; inner ramus of uro-
pod 2 shortened, partially fused to peduncle. Uro-
pod 3 small, rami in female not longer than pedun-
cle, article 2 of outer ramus very small, carrying 2
long apical setae. Telson ordinary to short, with
only one apical spine on each lobe plus setules.
DESCRIPTION.?Rostrum constricted, small. An-
tenna 1 especially thin; fuzz on article 1 of antenna
1 in male present; calceoli on male primary flagel-
lum of antenna 1 present. Calceoli on article 5 of
male antenna 2 present; flagellum in male with cal-
ceoli. Prebuccal parts strongly bulging forward, dis-
tinct, neither epistome nor upper lip dominant.
Right lacinia mobilis bifid, thin; mandibular palp
thin, article 1 slightly elongate, article 2 with outer
NUMBER 245 155
setae, apex of article 3 oblique. Lower lip bearing
cones. Outer plate of maxilla 1 with 11 spines, one
spine especially thickened. Inner plates of maxil-
liped thick, ordinarily setose. Coxae 2-3 elongate,
coxae 2-4 without special anterodorsal humps, one
ventral seta on coxae 1-3 especially thick. All pos-
terior spines on article 6 of pereopods 1-2 thick and
stiff, midapical seta present. Article 2 of pereopod 5
without facial setae. Peduncle of uropod 1 with
dorsolateral spines confined apically, medial spines
widely spread; peduncle of uropod 2 with only one
medial spine confined apically. Peduncle of uropod
3 lacking extra subapical setae or spines. Telson
with ordinary pair of midlateral or dorsal setules
on each side.
TYPE-SPECIES.?Matong matong, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?Matong differs from Ponthar-
pinia in the elongate article 2 of antenna 1, the
presence of cones on the lower lip, the reduction of
setules on the face of article 3 on antenna 2 to the
normal 2 in number, the normal fully biramous
right lacinia mobilis, the nontriturative molar, the
presence of posteroproximal setae on article 5 of
pereopods 1-2, and the shortened rami of uropod
3. In addition, Matong differs in these secondary
characters: absence of dorsal setae on article 4 of
antenna 2, enlarged article 5 of antenna 2, thin
mandibular palp, reduction of setae to 4 on inner
plate of maxilla 1, development of 2 stout spines
on the inner plate of the maxilliped, fully discrete
nail on dactyl of maxilliped, apical confinement
of dorsolateral spines on peduncle of uropod 1, loss
of supernumerary setae on dactyl of pereopod 5,
on peduncle of uropod 3, on article 2 of pereopods
3-4, on the telson, on urosomite 1, and the normal
phoxocephalid telson with apical spines and dorso-
lateral pairs of setules.
Matong differs from Mandibulophoxus and
Basuto in the elongate article 2 of antenna 1, the
absence of a special spine on the peduncle of uro-
pod 1, the well developed molars, the elongate wrist
of gnathopod 2, and the short rami of uropod 3.
Matong has almost reached the evolutionary
grade of Birubius except that it retains enlarged
gnathopods. Matong differs from Birubius also in
the shortened rami of uropod 3, the rudimentary
fusion of the inner ramus of uropod 2 to the pedun-
cle, the more apically placed ventral setae on article
2 of antenna 1, and the slightly developed, but clearly
present, posterior setae on epimera 1-2.
Matong matong, new species
FIGURES 61-64
Paraphoxus sp., Australia, J. L. Barnard, 1960:285-286.
DESCRIPTION OF FEMALE.?Head about 17 percent
of total body length, greatest width about 90 per-
cent of length; rostrum almost unconstricted, broad,
short, exceeding apex of article 1 on antenna 1.
Eyes medium, clear of pigment. Article 1 on pedun-
cle of antenna 1 about 1.4 times as long as wide,
about 2.3 times as wide as article 2, ventral margin
with about 6 setules and spinule, unproduced
dorsal apex with 3 setules; article 2 about 0.95
times as long as article 1, with apicoventral cycle of
4 setae; primary flagellum with 9 articles, about
0.75 times as long as peduncle, aesthetascs poorly
developed; accessory flagellum short, with 5 arti-
cles. Spine formula on article 4 of antenna 2 = 1-3-
4-3 or 1-3-4-4, dorsal margin without notch, bear-
ing one seta, ventral margin with 6 groups of 1-2
long to medium setae, one ventrodistal long spine;
article 5 about 0.8 times as long as article 4, facial
spine formula = 1-2-2-1, dorsal margin naked,
ventral margin with 4 sets of 1-2 long to short seta-
spines, 3 ventrodistal long spine-setae; flagellum
about 0.75 times as long as articles 4-5 of peduncle
combined, with 9 articles. Mandibles with weak
palpar hump; right incisor with 3 teeth; left incisor
with 5 humps in 2 branches; right lacinia mobilis
bifid, distal branch much shorter than proximal,
distal branch narrow, simple, pointed, proximal
branch simple, pointed, with marginal denticles;
left lacinia mobilis with 4 teeth; right rakers 9 plus
2 rudimentaries; left rakers 10; molar in form of
elongate plaque, right molar with 6 long to me-
dium spines, two short spines weakly disjunct, left
molar with 4 long to short spines, none disjunct,
each molar with plume; palp article 1 short, article
2 with one long inner apical seta and 3 other long
and short inner setae, plus one outer seta, article 3
almost 0.95 times as long as article 2, oblique apex
with 7-8 spine-setae, basofacial formula = 2-2 or
1-2. Inner plate of maxilla 1 enlarged, bearing one
long apical pluseta, one shorter similar apicomedial
seta, 1-2 apicolateral much shorter setae; palp arti-
cle 2 with 6 apical-medial marginal spines and
setae. Plates of maxilla 2 extending equally, outer
156 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
bS
FIGURE 61.?Matong matong, new species, holotype, female "a," 6.50 mm (b ? female "b," 6.45
mm; c ? male "c," 4.80 mm; v = male "v," 4.70 mm).
NUMBER 245
157
FIGURE 62.?Matong matong, new species, holotype, female "a," 6.50 mm (v ? male "v," 4.70 mm).
158 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
cRS
FIGURE 63.?Matong matong, new species, holotype, female "a," 650 mm (b ? female "b," 6.45
mm; c ? male "c" 4.80 mm; / ? female "f,f 6.80 mm; v ? male "v," 4.70 mm).
slightly narrower than inner, with 2 apicolateral
setae, inner with one medial seta. Inner plate of
maxilliped with 2 large, thick apical spines, 3 apico-
facial setae, 5 medial setae; outer plate with 11 +
medial and apical spines, 3 apicolateral setal spines;
palp article 1 with apicolateral seta, article 2 with
one apicolateral seta, medial margin of article 2
moderately setose, article 3 unprotuberant, with 4
facial setae, 2 lateral setae, nail of article 4 medium,
with 2 accessory setules. Coxa 1 strongly expanded
distally, anterior margin weakly concave; main ven-
tral setae of coxae 1?4 = 15-15-13-7, posteriormost
seta of coxae 1-3 shortened, one middle seta espe-
cially stout; anterior and posterior margins of coxa
4 strongly divergent, posterior margin very oblique,
almost straight, posterodorsal corner rounded, pos-
terodorsal margin short, almost straight, width-
length ratio of coxa 4 = 16:19. Long posterior setae
on article 2 of gnathopods 1-2 and pereopods 1-2 =
4-5-4-5, long anteriors = 1-6-0-0, short anteriors
4-3-5-2, no others. Gnathopods enlarged, hands
quadriform, gnathopod 1 sub-eusirid, article 3
slightly elongate on both pairs; width ratios of arti-
cles 5-6 on gnathopods 1-2 = 28:50 and 31:58, length
ratios = 62:66 and 61:74, palmar humps very large,
palms transverse; article 5 of gnathopod 1 ovate,
posterior margin rounded-flat; article 5 of gnatho-
pod 2 ovate, posterior margin rounded, short.
Pereopods 1-2 similar, facial setae formula on arti-
cle 4 = 5-6 and 5, on article 5 = 4 and 6, main
spine of article 5 extending to M. 75 on article 6,
article 5 lacking proximoposterior spines; spine
formula of article 6 = 3 + 4 and 4 + 4 plus mid-
distal seta, acclivity on inner margin of dactyls of
pereopods 1-2 sharp, produced as tooth, emergent
setule short, midfacial plusetae very long. Coxae 5-7
NUMBER 245
FIGURE 64.?Matong matong, new species, holotype, female "a," 6.50 mm (6
mm; c ? male "c," 4.80 mm; v ? male "v," 4.70 mm).
female "b," 6.45
posteroventral seta formula = 6-4-8. Articles 4-5 of
pereopods 3-4 of ordinary width, facial spine rows
moderately developed, facial ridge formula on arti-
cle 2 of pereopods 3-5 = 1-2-2, width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 45:40:34:16, of
pereopod 4 = 53:31:27:11, of pereopod 5 = 70:21:
22:7, length ratios of pereopod 3 = 68:34:33:39, of
pereopod 4 = 80:54:40:47, of pereopod 5 = 100:26:
24:21; article 2 of pereopod 5 reaching apex of arti-
cle 4, most posterior of ventral setules often slightly
elongate; medial apex of article 6 finely combed,
bearing 3 digital processes. Posteroventral corner of
epimeron 1 subquadrate, weakly produced, poste-
rior margin weakly convex, setulose, corner with
seta, anteroventral margin with 5 long setae, pos-
teroventral margin with 2 long setae; posteroventral
corner of epimeron 2 rounded, quadrate, with pos-
terior margin weakly setulose, facial setae = 9-12,
fully spread, or occasional 1-2 pairs set vertically;
posteroventral corner of epimeron 3 weakly protu-
berant, with setule sinus and small tooth, posterior
margin weakly convex, with 2 setule notches and
160 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
posterodistal seta set in weak notch, ventral margin
with 8-9 setae evenly spread, face with 1-3 setae
near ventral margin in middle. Urosomite 1 with 3
ventral setules at base of uropod 1, row of 4 short
facial setae, articulation line complete; urosomite 3
protuberant dorsally. Rami of uropods 1-2 with
articulate enlarged apical nails, outer ramus of
uropod 1 with 4 dorsal spines, inner with one,
outer ramus of uropod 2 with 1-2 dorsal spines,
inner with one dorsomedial spine; peduncle of uro-
pod 1 with 4 apicolateral spines and 3-6 basofacial
short setae, medially with 4-5 marginal setae plus
apical short, stout spine; peduncle of uropod 2 with
4-5 dorsal spines, and 0-1 basofacial seta, medially
with one small apical spine. Peduncle of uropod 3
with one ventral setule, dorsally with lateral spine
and setule, 2 medial setules; rami feminine, inner
extending to M. 55 on article 1 of outer ramus,
apex with one seta, medial and lateral margins
naked, article 2 of outer ramus short, 0.09, bearing
2 long setae, apicomedial margin of article 1 with 2
setae, lateral margin with 4 acclivities, setal-spine
formula = 1-1-1-1-2, setal formula = 0. Telson
short, length-width ratio = 25:32, not fully cleft,
each apex wide, obliquely truncate, lateral acclivity
absent, bearing ordinary lateral setule, spine next
medial shorter than setule, midlateral setules di-
verse. Cuticle with bulbar setules on edges of clear
spaces in midst of dense polygonal structure, emer-
gent setules especially long.
OBSERVATIONS (female).?Armament formulas on
article 1 of outer ramus on uropod 3, female "g,"
setae = 1-1-1-1-2 or 1-2-2; female "f," setae =
1-1-3, spines = 0-1-0; uropod 1 with mediobasal
setae on peduncle.
DESCRIPTION OF MALE (male "v," 4.70 mm, dam-
aged).?Eyes enlarged, causing extreme lateral pro-
tuberance of head. Article 1 of antenna 1 with
medial patch of fuzz; primary flagellum with 9
articles, one calceolus each on articles 2-5; accessory
flagellum with 5 articles. Spine formula on article 4
of antenna 2 = 3-4-4, on article 5 = 2-2, dorsal
margin of article 5 with 3 sets of male setae and
2 calceoli, ventral apex with 2 thin spine-setae; flag-
ellar formula = ?, 2-4, 6, 8 . . . 26 (broken), or ?,
2-5, 7, 9 . . . 25 (broken). Ventral margin of upper
lip slightly more excavate than in female. Distal
branch of right lacinia mobilis broadened, sub-
bifid; rakers 8 on each side; right molar spines 8,
left 7, no plume on left; right basofacial palp arti-
cle 3 = 2-2, left = 2-3. Inner plate of maxilla 2
with 4 setae, 2 large, 2 small. Maxilla 2 with 3 me-
dial setae on inner plate; 3 apicolateral setae on
outer plate. Palp article 4 of maxilliped with only
one accessory setule. Cnathopods slightly thinner
than in female. Article 2 of only pereopod 5 nar-
rower than in female. Epimera 1-3 broadened but
not protuberant posteriorly; posterior margin of
epimeron 3 not shortened; setal formulas, epimeron
1 anteroventral = 5, posteroventral = 3, epimeron
2 facial = 7-8, epimeron 3 posterior = 2 setules,
ventral = 6. Urosomite i with 6 faciolateral setae,
no ventral setuies. Spine formulas of uropods, uro-
pod 1 peduncle apicolateral = 4, basofacial = 5,
longer than in female, uropod 2 peduncle dorsal =
5, basofacial = 1, dorsal spines on outer ramus of
uropod 1 = 3, of uropod 2 = 2, inner ramus of uro-
pod 1 = 1, of uropod 2 = 1 ; ventral spines on pe-
duncle of uropod 3 = 0, spine formula on article 1
of outer ramus = 1-1-0-1-0-1-0-0, setal formula =
1-1-1-1-1-1-1-2, article 2 of outer ramus vestigial.
Telson elongate, distal spines scarcely shortened,
thin, each lobe with supernumerary lateral setule
and row of dorsal denticles.
OBSERVATIONS (male).?Right epimeron 3 of male
"v" lacking ventral setae.
DESCRIPTION OF JUVENILE (juvenile female "h,"
4.86 mm).?Spine formula on article 4 of antenna
2 = 1-3-4-4, on article 5 = 1-2-2; spine formulas,
peduncle of uropod 1 = 3 , basofacial = 2, outer
ramus = 3, uropod 2 peduncle = 3. outer ramus =
1; setal formulas, epimeron 1 anteroventral = 2,
posteroventral = 2, epimeron 2 facial = 5, epi-
meron 3 ventral = 3, no facial; outer ramus of uro-
pod 1 with setal formula of 1-2 on article 1, inner
apex with one seta, apical seta on inner ramus very
short, only as long as article 2 of outer ramus (in-
cluding immersed part of article 2).
ILLUSTRATIONS.?Male "v", on dorsal view of
head rostrum not fully extended because of down-
ward curve; right eye damaged; epimeron recon-
structed, right epimera 2-3 apparently aberrant,
epimeron 2 extending greatly ventrad, epimeron 3
asetose, left epimera apparently normal as illus-
trated.
HOLOTYPE.?NMV, female "a," 6.50 mm.
TYPE-LOCALITY.?CPBS 32N/866, 25 Aug 1966,
Western Port, Victoria, Australia, 13.3 m, sandy
gravel.
VOUCHER MATERIAL.?CPBS C3/4: female "b,"
NUMBER 245 161
6.45 mm (illus.). CPBS A2/2: subaduk male "c,"
4.80 mm (illus.). PPBES 126: male "v," 4.70 mm
(illus.). PPBES 928/4: female "f," 6.80 mm (illus.);
female "g," 7.45 mm; juvenile female "h," 4.86 mm.
MATERIAL.?CPBS, 28 samples from 14 stations
(49); PPBES, 6 samples from 4 stations (8); WPBES,
one sample from one station (1); AM, one sample
("Type" of Birubius batei, type erroneous) (2).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 8-19.2 m, sand, muddy sand. New
South Wales: Port Jackson.
Kotla, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
elongate, ventral setae confined almost apically.
Article 1 of antenna 2 not ensiform; article 3 with
2 setules; facial spines on article 4 in 3 or more
rows; article 5 ordinary in size. Right mandibular
incisor with 3 teeth; molar not triturative, ordinary,
medium, pillow-shaped, bearing 4 or more splayed,
semiarticulate spines, usually bearing fuzz; palpar
hump small. Palp of maxilla 1 biarticulate; inner
plate with 4 setae. Setation of maxilla 2 ordinary.
Inner plate of maxillipeds ordinary; apex of palp
article 3 not strongly protuberant, dactyl elongate,
apical nail distinct, elongate. Gnathopods similar,
weakly to moderately enlarged; article 5 of gnatho-
pods free, elongate, with weak eusirid attachment;
palms almost transverse, hands of gnathopods broad-
ened, poorly setose anteriorly. Article 5 of pereo-
pods 1-2 with posteroproximal setae. Article 2 of
pereopod 3 of broad form, artticles 4-5 of pereopods
3-4 broad to medium, article 2 of pereopods 3-4
not setose posteriorly; pereopod 5 ordinary, but
articles 4-6 enlarged, article 2 naked ventrally ex-
cept for one elongate setule, article 3 ordinary,
dactyl normal. Epimera 1-2 bearing large vertical
facial brushes of setae; epimeron 3 ordinary. Uroso-
mite 1 generally naked except for sparse apicoventral
setae or spines near base of uropod 1; urosomite 3
without dorsal hook or special process. Pe-
duncle of uropod 1 normally elongate, without
apicoventral spike, without special enlarged apico-
lateral-medial spine; peduncular apices of uropods
1-2 not combed; inner ramus of uropod 1 with
marginal spines in one row, no rami continuously
spinose to apex, inner ramus of uropod 2 fused to
peduncle. Uropod 3 small, article 2 of outer ramus
vestigial, carrying 2 medium apical setae. Telson
ordinary, with only one apical spine plus setule,
without special dorsal and lateral spines or setae.
DESCRIPTION.?Rostrum constricted, obsolescent.
[Fuzz on article 1 of antenna 1 in male unknown,
calceoli on male antenna 1 and antenna 2 un-
known.] Prebuccal parts ordinary, distinct, upper
lip dominant. Right lacinia mobilis bifid, thin;
mandibular palp thin, article 1 slightly elongate,
article 2 without outer setae, apex of article 3
oblique. Lower lip bearing cones. Outer plate of
maxilla 1 with 11 spines, one spine especially thick-
ened. Inner plates of maxilliped ordinarily setose.
Coxae 1-4 very short, without special anterodorsal
humps. All posterior spines on article 6 of pereo-
pods 1-2 thick and stiff, midapical seta present.
Article 2 of pereopod 5 without facial setae. Pedun-
cle of uropod 1 with dorsolateral spines widely
spread, medial spines confined apically, peduncle
of uropod 2 with medial spine or setule absent.
Peduncle of uropod 3 lacking extra subapical setae
or spines. Telson with ordinary pair of midlateral
or dorsal setules on each side.
TYPE-SPECIES.?Kotla batturi, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This genus appears to be de-
scendant from ancestors near Matong, new genus.
The resemblance is strong in the following charac-
ters: thin article 2 of antenna 1 with ventral setae
shifted apically, short rami and elongate peduncle
of uropod 3, broadened hands of gnathopods,
stubby rami of uropod 2, and truncate apices of
telson. Kotla differs from Matong in the fully fused
inner ramus of uropod 2, the jewel-like spines on
uropod 2, the large vertical brushes of setae on
epimera 1-2, the short coxae, the almost fully
spread dorsolateral spination on the peduncle of
uropod 1, the loss of basomedial setation on the
peduncle of uropod 1 and the loss of major teeth on
the left lacinia mobilis.
Kotla batturi, new species
FIGURES 65-67
DESCRIPTION OF FEMALE.?Head about 15 percent
of total body length, greatest width about 100 per-
cent of length; rostrum obsolescent, scarcely reach-
ing along article 1 on antenna 1. Eyes large, stained
ochraceous, clear of occluding pigment. Article 1 on
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s
FIGURE 65.?Kotla batturi, new species, holotype, female "a," 5.10 mm (b ? female "b," 4.25 mm;
c ? male "c," 4.33 mm).
NUMBER 245 163
FIGURE 66.?Kotla batturi, new species, holotype, female "a," 5.10 mm.
164 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
peduncle of antenna 1 about 1.5 times as long as
wide, about twice as wide as article 2, ventral mar-
gin with about 9 setules, weakly produced dorsal
apex with 3?4 setules; article 2 about 1.1 times as
long as article 1, with ventral cycle of 3-4 setae near
apex; primary flagellum with 8 articles, about 0.85
times as long as peduncle, bearing aesthetascs; ac-
cessory flagellum with 4 articles. Spine formula on
article 4 of antenna 2 = 1-3-5-5-4 or 1-3-5-5-3 or
1-4-5-5-3, dorsal margin naked, ventral margin
with apical group of 4 medium setae, one ventro-
distal short spine and brush of large mediodorsal
setules; article 5 about 0.9 times as long as article 4,
facial spine formula = 2-2, dorsal margin bearing
setule, ventral margin with 3 sets of 1-3 long to
medium spines, 3 ventrodistal long setae, one sub-
distal facial spine; flagellum about 0.65 times as
long as articles 4-5 of peduncle combined, with 7
articles. Mandibles with weak palpar hump; right
incisor with 3 teeth; left incisor with 3 humps in 2
branches; right lacinia mobilis bifid, distal branch
much shorter than proximal, narrow, pointed, den-
ticulate, proximal branch simple, pointed, with
marginal denticles; left lacinia mobilis with obso-
lescent tooth notches; right rakers 6 plus one rudi-
mentary; left rakers 7 plus one rudimentary; molar
in form of elongate plaque, bulbous, right molar
with 8-9 long to medium spines, none disjunct, left
molar with 8 long to medium spines, one of these
weakly disjunct, each with plume; palp article 1
elongate, article 2 with pair of long to medium in-
ner apical setae and 3 other long and short inner
setae, article 3 about 0.8 times as long as article 2,
scarcely oblique apex with 6 spine-setae, basofacial
formula = 1-2. Inner plate of maxilla 1 ordinary,
bearing one long apical pluseta, one similar apico-
medial seta, 2 apicolateral much shorter setae; palp
article 2 with 6 apicalmedial setae. Plates of maxilla
2 extending subequally, outer much narrower than
inner, outer with 4 apicolateral setae, inner with 3
medial setae. Inner plate of maxilliped with 2 large
thick apical spines, 3 apicofacial setae, 4 medial setae;
outer plate with 7 medial and apical spines, 2 apico-
lateral setae, one apical cone; palp article 1 with 2
apicolateral setae, article 2 with 2 apicolateral setae,
medial margin of article 2 moderately setose, article
3 with 5 facial setae, 3 lateral setae, nail of article
4 long, with 3 accessory setules. Coxa 1 scarcely ex-
panded distally, anterior margin almost straight;
main ventral setae of coxae 1-4 = 3-4-5-3 or 3-3-
4-2, posteriormost seta of coxae 1-3 slightly short-
ened; anterior and posterior margins of coxa 4
strongly divergent, posterior margin very oblique,
straight, posterodorsal corner rounded, postero-
dorsal margin long, undulant, width-length ratio
of coxa 4 = 37:32. Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 = 7-10-(8-9)-
(10-11), short anteriors = l-O-(2-4)-3, long ante-
riors = 4(+ 3 facial)-3-0-0, short anteriors = (3-4)-
(3-4)-7-(8-9), no others. Gnathopods with stout
hands, article 3 elongate, width ratios of articles
5-6 on gnathopods 1-2 = 24:48 and 28:48, length
ratios = 75:72 and 75:71; palmar humps large,
palms weakly oblique; article 5 of gnathopods 1-2
elongate, ovate, posterior margin rounded. Pereo-
pods 1-2 similar; facial setae formula on article
4 = 4-5 and 5-6, on article 5 = 2-3 and 2-4; main
spine of article 5 extending to M. 80 on article 6,
article 5 with 2-3 proximoposterior spines; spine
formula of article 6 = 4 + 3 or 3 + 3 plus middistal
seta, most spines especially short, acclivity on inner
margin of dactyls of pereopods 1-2 sharp, produced
as tooth, emergent setule long, stout, midfacial
pluseta long, dactyls short and stout. Coxae 5-7
posteroventral setule formula = 2-2-7. Articles 4-5
of pereopods 3-4 of ordinary width, facial spine
rows medium to sparse respectively, facial ridge
formula on article 2 of pereopods 3-5 = 1-2-2, that
on pereopod 3 very short; width ratios of articles
2, 4, 5, 6 of pereopod 3 = 49:36:38:23, of pereo-
pod 4 = 58:34:32:16, of pereopod 5 = 75:29:28:13,
length ratios of pereopod 3 = 70:31:37:43, of pereo-
pod 4 = 90:55:52:50, of pereopod 5 = 108:36:33:43;
article 2 pereopod 5 reaching middle of article
4; medial apex of article 6 smooth. Posteroventral
corner of epimeron 1 rounded, posterior margin
almost straight, with setule, anteroventral mar-
gin with 4-9 short setae, posterior face with vertical
row of 11-15 long setae, ventralmost weakly dis-
junct; posteroventral corner of epimeron 2 weakly
protuberant, guarded by setule sinus, posterior mar-
gin weakly convex, with 2 setule notches, facial setae
= 3?5 in horizontal disarray plus upper vertical row
of 5-7 long setae, facial ridge absent; posteroventral
corner of epimeron 3 rounded, weakly protuberant,
with setule sinus, posterior margin convex, with
setule notches above and below corner, ventral face
with horizontal group of 5-7 setae in disarray near
ventral margin in middle. Urosomite 1 with one
lateral and 2 ventral setules at base of uropod 1,
NUMBER 245 165
FIGURE 67.?Kotla batturi, new species, holotype, female "a," 5.10 mm (c ?= male "c," 4.33 mm;
Y = flagellum of antenna 2).
articulation line complete; urosomite 3 strongly
protuberant dorsally. Rami of uropods 1-2 with
articulate enlarged apical nails, outer ramus of uro-
pod 1 with 5-6 dorsal spines, inner with 3, outer
ramus of uropod 2 with 2 dorsal spines, inner with
2 dorsal spines, one thin and one jewel like, all
other spines of uropod 2 jewel-like, inner ramus
fused to peduncle, uropod 2 very thick, peduncle of
uropod 1 with 14-15 dorsolateral spines and 3 short
basofacial setae, medially with 2 apical spines in
tandem, peduncle of uropod 2 with 4 dorsal spines,
medially naked (fused). Peduncle of uropod 3 with
one ventral setule, dorsally with one lateral spine,
one medial spine; rami not longer than peduncle,
submasculine, inner extending to M. 80 on article 1
of outer ramus, apex with 3 setae, medial margin
setose, article 2 of outer ramus short, 0.07, bearing 2
medium setae, medial margin of article 1 with 3
setae, lateral margin with 4 acclivities, spine form-
ula = 0, setal formula = 1-1-1-1-2. Telson ordi-
nary, length-width ratio =1:1 , not fully cleft, each
apex wide, truncate, lateral acclivity absent, bear-
ing long lateral setule, spinule or setule next medial
shorter, no spines, midlateral setules diverse. Cuti-
cle with bulbar setules mixed with 5-10 times as
many pipes, setules surrounded by cordate plaques,
166 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pipes emerging as hemicylinders, setules ordinary.
DESCRIPTION OF MALE (very young male "c," 4.33
mm).?Like female (but because of youth with
much smaller eyes). Antenna 1 similar to female, no
medial fuzz. Antenna 2 stouter, flagellum slightly
longer, articles thick, articles 4-5 with similar spine
formula. Setal formula of coxae 1-4 = 2-3-4?2.
Epimeron 1 with 6 anteroventral setae and 12 setae
in posterior brush (one seta disjunct); epimeron 2
with 6 setae in brush and 3 horizontal setae in
tandem; epimeron 3 slightly elongate anteroposte-
riorly, with 4 setae in tandem, posteroventral corner
with 3 small teeth and 4 setule-notches. Uropod 1
with 3 basofacial setae, 16 peduncular spines, outer
ramus with 6 spines, inner ramus with 2; uropod
2 like female holotype. Inner ramus of uropod 3
elongate, as long as outer ramus but poorly setose,
5 lateral setae of outer ramus not plumose but with
short scales.
OBSERVATIONS.?Ventral setules on article 4 of
antenna 2 large and placed more distally than
normal; accessory setules between rakers on man-
dibles absent; dactyls of gnathopods apically flagel-
late. The absence of facial setae on article 5 of
pereopod 1 and the presence of an extra pair of
posterior spines are aberrations in the holotype.
The holotype female is almost senilic in light of
these aberrations and in the view of the well devel-
oped eyes and head; see lateral view of male head
also typical of normal females.
ILLUSTRATIONS.?Dorsal view of holotype head
not illustrated because of damage; thick apical spines
on inner plate of maxilliped shown in Figure 66: Si-
HOLOTYPE.?NMV, female "a," 5.10 mm.
TYPE-LOCALITY.?CPBS 600/5, 29 Mar 1965,
Western Port, Victoria, Australia, 18.3 m, sand, silt.
VOUCHER MATERIAL.?CPBS 600/3; female "b,"
4.25 mm (illus.); male "c," 4.33 mm (illus.); female
"d," 4.80 mm.
MATERIAL.?CPBS, 3 samples from 2 stations
(10).
DISTRIBUTION.?Victoria, Western Port, 18.3 nt,
sand, silt.
Yammacoona, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
ordinary to elongate, ventral setae widely spread.
Article 1 of antenna 2 not ensiform; article 3 with
2 setules; facial spines on article 4 in 2 rows; article
5 short. Right mandibular incisor with 3 teeth;
molar not triturative, ordinary, pillow-shaped, bear-
ing 4 or more splayed, semiarticulate spines, bear-
ing fuzz; palpar hump medium to large. Palp of
maxilla 1 biarticulate; inner plate with 4 setae.
Setation of maxilla 2 ordinary. Inner plate of max-
illipeds ordinary, apex of palp article 3 moderately
protuberant, dactyl elongate, apical nail distinct,
medium. Gnathopods dissimilar, gnathopod 2 mod-
erately enlarged; article 5 of gnathopods 1-2 short,
but free, without eusirid attachment; palms oblique,
hand of gnathopod 2 broadened, poorly setose ante-
riorly; article 5 of pereopods 1-2 with posteroproxi-
mal spines. Article 2 of pereopod 3 of broad form,
articles 4-5 of percopods 3-4 broad, article 2 of
pereopods 3-4 not sciosc posteriorly; pereopod 5
ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Kpimera 1-2 lacking numerous
long posterior setae, without midfacial setae above
ventral facial ridge; epimeron 3 ordinary. Uroso-
mite 1 generally naked except for sparse apicoven-
tral setae or spines near base of uropod 1; uroso-
mite 3 without dorsal hook or special process. Pe-
duncle of uropod 1 normally elongate, without
apicoventral spike, without special enlarged apico-
lateral-medial spine; peduncular apices of uropods
1-2 not combed; spine(s) on inner ramus of uropod
1 in one row, some rami with accessory nails, inner
ramus of uropod 2 ordinary. Uropod 3 ordinary,
article 2 of outer ramus carrying 2-3 medium to
long apical setae. Telson ordinary, with only one
apical spine plus setae on each lobe plus setules.
DESCRIPTION.?Rostrum constricted, obsolescent.
Fuzz on article 1 of antenna 1 in male present; cal-
ceoli on male primary flagellum of antenna 1 pres-
ent. Calceoli on article 5 of male antenna 2 present;
flagellum in male with calceoli. Prebuccal parts
strongly extended forward, almost horizontal, espe-
cially massive, poorly separated from each other,
upper lip dominant. Right lacinia mobilis bifid,
flabellate; mandibular palp thin, article 1 highly
elongate, article 2 without outer setae, apex of
article 3 transverse. Lower lip bearing cones. Outer
plate of maxilla 1 with 11 spines, one spine espe-
cially thickened. Inner plates of maxilliped espe-
cially thin, poorly armed; outer plates small. Coxae
2-A without special anterodorsal humps. All pos-
terior spines on article 6 of pereopods 1-2 thick and
stiff, midapical seta absent. Article 2 of pereopod 5
NUMBER 245 167
without facial setae. Peduncle of uropod 1 with
dorsolateral spines confined apically, medial spines
confined apically; peduncle of uropod 2 with only
one medial setule confined apically. Peduncle of
uropod 3 lacking extra subapical setae or spines.
Telson with ordinary pair of midlateral or dorsal
setules on each side.
TYPE-SPECIES.?Yammacoona kunarella, new spe-
cies.
COMPOSITION.?Unique.
RELATIONSHIP.?Yammacoona appears to have its
closest relationship to Matong and Kotla but differs
from those genera in many significant characters, as
follows: stouter antenna I with shorter article 3 and
with the ventral setae of article 2 placed proximally;
reduction in number of spine rows on article 4 of
antenna 2, with a longer main row; enlarged palpar
hump on the mandible; smaller plates of the maxil-
lipeds and the protuberant article 3 of the palps;
free inner ramus of uropod 2; and, to a greater or
lesser extent, the shortened article 5 of gnathopod
2. Yammacoona furthermore differs from Kotla in
the absence of vertical setal brushes on the faces of
epimera 1-2 but does bear such a brush on epi-
meron 3; and it differs from Kotla in the narrow
apical confinement of the dorsolateral spines on the
peduncle of uropod 1 and in the enlarged articles
4-5 of pereopod 5. Though bifid, the right lacinia
mobilis is far closer to that of Pontharpinia, lacking
that proximal part possibly representing a fused
raker spine seen in Matong and Kotla.
The enlarged palpar hump on the mandible sug-
gests that Yammacoona belongs to the Pontharpinia-
Mandibulophoxus-Basuto group of genera. The
closest similarity seems to occur between Basuto
and Yammacoona. A conspicuous resemblance be-
tween the two genera occurs in epimeron 3, gna-
thopod 2 and the short article 5 of antenna 2, but
Yammacoona differs in the broadness of article 2
on pereopod 3, the unusual head, smaller mandibu-
lar palp, small outer plate of maxilliped, unpro-
duced article 3 of the maxillipedal palp, longer
article 2 of antenna 1, completely distinctive coxa 4,
and the short article 2 on the outer ramus of uro-
pod 3. Yammacoona is even more remote from
Mandibulophoxus than from Basuto.
Yammacoona resembles Pontharpinia in the
palpar hump of the mandible, thin articles 4-5 of
pereopod 5, short article 5 of antenna 2 and in the
gnathopods, but differs in many fundamental char-
acters, among them the simple molar, small outer
plate of the maxilliped, weaker mandibular palp,
shape of epimeron 3 and coxae 1-4, short article 2
on outer ramus of uropod 3, longer article 2 of an-
tenna 1, shape of the head, nonsetose dactyl of
pereopod 5, and absence of continuous spination
on the rami of uropods 1?2.
Although gnathopod 2 is enlarged, the wrists of
the gnathopods are shortened, article 5 of antenna
2 is shortened and the margins of epimeron 3 are
relatively simple. Yammacoona otherwise has few
similarities to the Brolgus group of genera and is
probably not a product of that group. Some of the
differences seen in Yammacoona are the short coxae,
distinctive article 2 of antenna 1, large palpar
hump on the mandible, absence of combs on uro-
pods 1-2, dissimilarities in pereopods 3-5 and inner
plate of maxilla 1, small outer plate of the maxil-
liped, and the presence of accessory apical nails on
uropods 1-2.
Among species of Birubiinae, Yammacoona has
the closest resemblance to K. cadgeeus in terms of
epimera, uropods 1-2, head, right lacinia mobilis,
shape of coxa 4, proximal placement of setae on
article 2 of antenna 1, thin articles 4-5 of pereopod
5 and protuberant article 3 of the maxilliped. But
Yammacoona differs from K. cadgeeus in the short-
ened and poorly setose coxae, enlarged gnathopod
2 with short wrist, and in the absence of a hook on
urosomite 3.
Yammacoona kunarella, new species
FIGURES 68-71
DESCRIPTION OF FEMALE.?Head about 12 percent
of total body length, greatest width about 75 per-
cent of length; rostrum unconstricted, broad, short,
antenna 1 attached directly to apex of rostrum.
Eyes large, mostly occluded with pigment. Article 1
on peduncle of antenna 1 about 1.45 times as long
as wide, about 1.4 times as wide as article 2, ventral
margin with 3 apical setules, weakly produced
dorsal apex with one setule; article 2 about 0.6
times as long as article 1, with ventral cluster of
3-6 setae on hump; primary flagellum with 9-11
articles, about 0.8 times as long as peduncle, bear-
ing aesthetascs; accessory flagellum with 9-11 arti-
cles. Spine formula on article 4 of antenna 2 = 1-
3-9 or 1-3-8 or 1-2-9, dorsal margin with notch
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 68.?Yammacoona kunarella, new species, holotype, female "a," 3.46 mm (v ? male "v,
3.55 mm; stippled area of vCn = mandible).
NUMBER 245 169
bearing 2 spine-setae, ventral marign with 3 me-
dium setae and one spine, one ventrodistal medium
spine; article 5 about 0.7 times as long as article 4,
facial spine formula = 5 or 4, dorsal margin naked,
ventral margin with 2 sets of 2 long and short setae,
3 ventrodistal short to medium spines; flagellum
about 1.4 times as long as articles 4-5 of peduncle
combined, with 9-11 articles. Mandibles with strong
palpar hump; right incisor with 3 teeth; left incisor
with 5 humps in 2 branches; right lacinia mobilis
bifid, distal branch little shorter than proximal,
distal branch flabellate, denticulate, proximal
branch flabellate, with marginal denticles and facial
humps; left lacinia mobilis with 5 teeth; right rak-
ers 7 plus one rudimentary; left rakers 7 plus 2-3
rudimentaries; molar in form of elongate plaque
bearing bulbous fuzzy hump, right molar with 5
medium to short spines plus one short spine strongly
disjunct, left molar with 3 medium spines plus one
short spine strongly disjunct, each molar with
plume; palp article 1 elongate, article 2 with one
short inner apical seta and one other shorter inner
seta, article 3 about 0.85 times as long as article 2,
truncate apex with 6 spine-setae, middorsal form-
ula = 2 spines. Inner plate of maxilla 1 enlarged,
bearing one long apical pluseta, one shorter apico-
medial seta, 2 much shorter apicolateral setae; palp
short and stout, palp article 2 with 5 long apical
and subapical setae. Plates of maxilla 2 extending
subequally, of equal breadth, outer with 2 apico-
lateral setae, inner with one medial seta. Inner
plate of maxilliped with one apical spine, 2?3
apicofacial setae, 2 medial setae; outer plate very
small and almost hidden behind inner plate, with
3-4 medial and apical spines, one apicolateral seta;
palp article 1 with apicolateral seta, article 2 elon-
gate, with one apicolateral seta, medial margin of
article 2 weakly setose, article 3 strongly protuber-
ant, with one facial seta, one lateral seta, nail of
article 4 medium, with 2 accessory setules. Coxae
1-4 short and broad, coxa 1 strongly expanded
apically, anterior margin straight; main ventral
setae of coxae 1-4 = 1-1-1-0; anterior and posterior
margins of coxa 4 strongly divergent, posterior mar-
gin very oblique, almost straight, posterodorsal
corner rounded, posterodorsal margin short, convex,
width-length ratio of coxa 4 = 1:1; long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1_2 = (2-3)-(2-3)-2-3, short anteriors = (0-l)-(2-
3)-(3-6)-(4-5), long anteriors = 0-0-1-0, short an-
teriors = 3-3-5-3, no others. Gnathopod 2 enlarged,
hand of gnathopod 1 slightly enlarged; width ratios
of articles 5-6 on gnathopods 1-2 = 23:39 and
22:50, length ratios = 47:62 and 34:80; palmar
humps very large, palms strongly oblique; article 5
of gnathopod 1 triangular, posterior margin lobate;
article 5 of gnathopod 2 short, triangular, lobate,
cryptic. Pereopods 1-2 similar; posterior setae on
pereopod 1 article 4 mainly apical, on pereopod 2
extending apically to mid mark, in groups 2-2-2-1;
facial setae formula on article 4 = 3 and 3, on arti-
cle 5 = 1 and 2; main spine of article 5 extending
to M. 100+ on article 6, article 5 with 3-4 proximo-
posterior spines; spine formula of article 6 = 3 + 3,
no middistal seta, most spines long; acclivity on in-
ner margin of dactyls of pereopods 1-2 sharp, pro-
duced as tooth, emergent setule long, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule
formula = 3-5-5. Articles 4-5 of pereopods 3-4
broad, facial spine rows dense, facial ridge formula
on article 2 of pereopods 3-5 = 0-1-2; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 50:50:50:22, of
pereopod 4 = 67:58:50:20, of pereopod 5 = 73:18:
17:10, length ratios of pereopod 3 = 78:43:38:45, of
pereopod 4 = 85:52:42:48, of pereopod 5 = 105:24:
24:30; article 2 of pereopod 5 exceeding apex of
article 5, dactyl weak, medial apex of article 6
smooth, posteroventral corner of epimeron 1
rounded, protuberant, posterior margin weakly si-
nuous, corner with setule and second setule above,
anteroventral margin with 3 medium setae, postero-
ventral face with 1-2 long setae in vertical pair;
posteroventral corner of epimeron 2 rounded,
weakly protuberant, guarded by setule sinus, poste-
rior margin weakly convex, facial setae = 5, poste-
riormost pair or triad set vertically, setae crowded
forward; posteroventral corner of epimeron 3
rounded, weakly protuberant, with setule sinus,
posterior margin almost straight, face with oblique,
almost vertical row of 3-6 setae. Urosomite 1 with
ventral setule at base of uropod 1, articulation line
complete; urosomite 3 scarcely protuberant dor-
sally. Rami of uropods 1-2 with articulate
enlarged apical nails, inner rami of uropods 1-2
with one accessory nail, outer ramus of uropod 1
with 1-2 dorsal spines, inner with one, outer ramus
of uropod 2 with one dorsal spine, inner with no
dorsomedial spine; peduncle of uropod 1 with one
apicolateral spine and microscopic basofacial setule,
medially with marginal setule and apical spine;
170 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
vES
FIGURE 69.?Yammacoona kunarella, new species, holotype, female "a," 3.46 mm (v ? male "v,"
3.55 mm).
NUMBER 245 171
peduncle of uropod 2 with 4-7 dorsal spines, me-
dially with one apical setule; apicolateral corners
of peduncles on uropods 1-2 lacking comb. Pedun-
cle of uropod 3 with 2 ventral spines, dorsally with
one lateral setule, one medial spine; rami feminine,
inner extending to M. 65 or less on article 1 of
outer ramus, apex with one long seta, medial and
lateral margins naked, article 2 of outer ramus
short, 0.17, bearing 3 long setae, medial margin of
article 1 naked, lateral margin with 3 acclivities,
spine formula = 2-2-2-2, setal formula = 0. Telson
short and broad, length-width ratio = 7:8, not fully
cleft, each apex wide, rounded-truncate, lateral ac-
clivity absent, bearing short lateral setule, spine
next medial much longer than setule, set sub-
apically, with adjacent very short medial setule,
midlateral setules diverse. Cuticle with ordinary bul-
bar setules, emergent setules especially short.
OBSERVATIONS (female).?Lateral cephalic lobe
thick, anteriorly truncate, strongly extended for-
ward. Apex of each flagellum with short aesthetasc
not counted as article. Mouthparts placed highly
ventrad, prebuccal complex almost horizontal,
strongly extended forward. Molar and spines
crowded towards incisor. Article 2 of pereopods 1-2
with additional small posterior straw-shaped setae
paired with long members and extending 3-4 in
number above long setae. Anterior margin of epi-
meron 1 with 1-10+ straw-like setules (highly vari-
able), ventral margin with 3-7 straw-like setules;
setae on face of epimeron 2 variable, additional
variations as follows: 3 vertical and 2 horizontal,
or 2 horizontal and 2 vertical, or 4 vertical and one
thrust forward horizonally. Basofacial setule on uro-
pod 1 so small as to be almost invisible.
DESCRIPTION OF MALE.?Head exceptionally elon-
gate, up to 21 percent of total body length, eyes
enlarged, anterior cephalic extension marked off
by ventral constriction. Articles 1-2 of antenna 1
of subequal width, article 1 relatively narrower
than in female, bearing medioventral patch of
long fuzz, article 2 not as strongly humped ven-
trally as in female, bearing 5 posterior setae in
cycle; primary flagellum with 11 articles, terminal
members bearing aesthetascs, one calceolus on each
of articles 1-7; accessory flagellum 8-9 articulate.
Article 3 of antenna 2 with only 2 small facial
setules, articles 4-5 greatly broadened, spines short
and broad, spine formula of article 4 = 1-3-9, of
article 5 = 7, dorsomedial faces of articles 3-4 with
setular fuzz, dorsal margin of article 5 with one
calceolus, ventral margin of article 4 with 2 club
spines and 2 setules, ventral margin of article 5
with one acclivity bearing spine and setule, ventro-
distal margin with 2 spines, seta and setule; flagel-
lum highly elongate, with 30 or 35 articles, one cal-
ceolus each on alternate articles, either odd or even,
including penultimate article, basally calceoli irre-
gular, for example on articles 3, 4, 6, 8 . . . or
3, 6, 8, 10 . . . or 3, 5, 7, 9 Raker row of
right mandible with 5 spines; left with 4 + 2
rudimentaries; right and left molars with 4
spines + one disjunct; left lacinia mobilis with
4 teeth and supernumeraries; palps with arti-
cles 1-3 more elongate than in female, article 1
about 65 percent as long as article 2, latter with 2
small and one large inner setae, article 3 with 2
outer acclivities bearing 1 and 3 spine-setae. Palp
of maxilla 1 with 4 apical setae. Plates of maxilla 2
much thinner than in female. Inner plate of maxil-
liped with only 2 apical setae. Single longer seta on
each of coxae 1-3 shorter than in female, coxae
much shorter than in female, broader than tall (see
illustrations). Long posterior setae of article 2 on
gnathopods 1-2 = (2)?(1?2), posterior margin also
bearing 6-8 stiff straw-like setae; articles 5-6 of
gnathopods of much different proportions from fe-
male, both gnathopods with broad article 6 and
elongate article 5, length ratios of articles 5-6 of
gnathopods 1-2 = 53:68 and 47:68, width ratios =
26:47 and 22:51; article 5 of gnathopod 1 with
medium-broad posterior lobe, of gnathopod 2 with
narrow protuberant lobe, article 5 almost cryptic
on gnathopod 2. Long posterior setae of article 2 on
pereopods 1-2 = 2 and 2 with 10-11 stiff and short
straw-like setae lining margin, some of these paired
with long setae or occasionally short setae paired
together, setal formula of article 4 of pereopods
1-2 = 3 and 3, of article 5 = 1 and (0 or rudimen-
tary), article 6 with rows of 3 and 3 spines, anterior
margin of article 2 on pereopods 1-2 with up to
18 straw-like setae. Coxa 5 slightly smaller and coxa
6 much smaller than in female; posteroventral
setule formula on coxae 5-7 = l-2-(3^i). Article 6
of pereopods 3-4 more elongate than in female,
articles 4-5 narrower, article 2 of pereopod 4 nar-
rower, spine densities much lower; pereopod 5 rela-
tively more elongate and/or thinner than in fe-
male, dactyl reduced, anterior margins of article 2
on pereopods 3-5 with dense fuzz composed of stiff,
172 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 70.?Yammacoona kunarella, new species, holotype, female "a," 3.46 mm (v ? male "v,1
3.55 mm).
NUMBER 245 173
FIGURE 71.?Yammacoona kunarella, new species, holotype, female "a," 3.46 mm (n ? female "n,1
4.20 mm; v -> male "v," 3.55 mm; w ? female "w," 4.0 mm; y ? male "y," 3.60 mm).
174 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
straw-like setules, article 2 of pereopod 4 with only
short anterior ridge. Epimera elongate anteroposte-
riorly, epimeron 1 without distinct ridge (illustra-
tion showing 2 artificial folds), with one antero-
ventral seta and posterior face with vertical row of
2-3 setae, posterior margin strongly bulbous, straw-
like setae absent; epimeron 2 with bulbous poste-
rior margin, no lateral ridge, or remnant of ridge,
row of 6 setae in weak crescent or 3 horizontal and
one highly disjunct seta or in groups of 2-1-1;
epimeron 3 with 2 or 3 facial setae set highly ven-
trad. Basofacial rudimentary setae of uropod 1 some-
what more distinct than in female, outer ramus
with 3-4 dorsal spines, inner with one; peduncle of
uropod 2 with 3-6 spines, apicomedial corner with
only pair of setules, outer ramus with 2 spines, in-
ner ramus of uropod 2 with basomedial spine. Uro-
pod 3 similar to that of terminal female but lack-
ing ventral peduncular spines, dorsally with one
lateral, one medial setule; rami of proportions
similar to terminal female, inner more strongly
setose, spines on acclivities of outer ramus con-
verted to setae, setal formula = 2-2-2-2-2. Telson
less cleft than in female, main apical spine on each
lobe very short, each lobe with mediodorsal row of
denticles.
DESCRIPTION OF JUVENILE (juvenile "u," 2.65 mm).
?Generally similar to adult female, but eyes very
small; flagella of antenna 1 with 11 primary and 9
accessory articles; spine formula of article 4 on
antenna 2 = 1-3-7, article 5 with 4 facial spines;
epimeron 1 with one long anteroventral seta, 4
straw-like anterior setules, 2 posterofacial setae (ver-
tical row), epimeron 2 with 3 facial setae (2 hori-
zontal, one thrust), epimeron 3 with 3 setae; uropod
1 peduncle with one spine, outer and inner rami
each with one dorsal spine, peduncle of uropod 2
with 4-5 spines, outer ramus with one, inner lack-
ing dorsal spines; uropod 3 with short inner ramus
(as illustrated for large female "n").
VARIATIONS.?Female holotype with straw-like
setae poorly developed in comparison with other
specimens; other females usually of larger body size
but female holotype generally with most advanced
morphology except in regard to straw-like setae of
epimeron 1. Specimens bearing glue-like exudate
creating great difficulty in dissecting and mounting
parts. Large female "n," 4.20 mm, with juvenoid
uropod 3 bearing short inner ramus (see illustra-
tion).
ILLUSTRATIONS.?Female: Antenna 1 drawn from
medial aspect but description based on observed
lateral aspect (as usual); outer lobule of article 2
on antenna 2 not adequately illustrated, poorly
preserved or often glued to article 3 by preserva-
tional exudate; lower lip of female "n" bearing off-
set flattened outer lobe; coxae in poor condition,
illustrations representing reconstructions; dactyl of
maxilliped bearing outer basal short pluseta hidden
by other structures; illustration of left pereopod 1
of holotype showing dotted seta on article 5, this
seta represented on right pereopod 1 but formed as
short spine on left pereopod 1 as shown.
Male: Sketch of head in Figure 68: vCn with an-
tenna 2 removed to show mandible (shaded), re-
mainder of structure being prebuccal complex (note
horizontal position of this structure); pereopod 3
drawn about 3 percent smaller than pereopod 5,
pereopod 4 drawn about 10 percent smaller than
pereopod 5; pereopods 3-5 larger relative to body
than in female.
HOLOTYPE.?NMV, female "a," 3.46 mm.
TYPE-LOCALITY.?CPBS 32/E2, 19 Feb 1965,
Western Port, Victoria, Australia, 12.8 m, sand.
VOUCHER MATERIAL.?Type-locality: female "b,"
3.34 mm. RHM: male "v," 3.55 mm (illus.); male
"y," 3.60 mm <illus.). CPBS 51N: female "w," 4.0
mm (in 2 pieces, illus). PPBES 979/1: female "n,"
4.20 mm (illus.); juvenile "u," 2.65 mm.
MATERIAL.?CPBS, 8 samples from 5 stations (11);
RHM, 2 samples from 2 stations (4); PPBES, 3
samples from 2 stations (4); WPBES, 2 samples
from 2 stations (3).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, neritic and 12.0-16.5 m, sand, muddy
sand.
PARHARPINI1NAE
DIAGNOSIS.?Article 2 of antenna 1 ordinary;
mandibular molar reduced to a small hump with
articulate spines; palp of maxilla 1 biarticulate;
setation on maxilla 2 ordinary; gnathopod 2 as
small as gnathopod 1; article 2 of pereopod 3 of
broad form, tapering distally; pereopod 5 ordinary.
SPECIAL CHARACTERS.?Telson with supernumer-
ary dorsal setae and spines; peduncle of uropod 1
with special apical spine.
DESCRIPTION.?Article 5 of antenna 2 of normal
size; epimeron 3 of nonrounded classification;
NUMBER 245 175
apices of peduncles on uropods 1-2 combed or not
combed.
TYPE GENUS.?Parharpinia Stebbing.
COMPOSITION.?Protophoxus K. H. Barnard.
Parharpinia Stebbing
Parharpinia Stebbing, 1899:207; 1906:147.?Barnard and
Drummond, 1976:531.
DIAGNOSIS.?Eyes present. Flagella of antennae 1-2
unreduced in female. Article 2 of antenna 1 ordi-
nary, ventral setae widely spread. Article 1 of an-
tenna 2 not or weakly ensiform; article 3 with 2
setules; facial spines on article 4 primarily in one
main row; article 5 ordinary in size. Right mandi-
bular incisor with 3 teeth; molar not triturative,
small, bearing 4 or more splayed semiarticulate
spines, usually bearing fuzz; palpar hump medium.
Palp of maxilla 1 biarticulate; inner plate with 4
setae. Setation of maxilla 2 ordinary. Inner plate of
maxillipeds ordinary; apex of palp article 3 weakly
protuberant, dactyl elongate, apical nail distinct,
mostly immersed, short. Gnathopods ordinary,
small, similar; article 5 of gnathopods 1-2 of ordi-
nary length, free, without eusirid attachment; palms
oblique, hands of gnathopods 1-2 ordinary, ovato-
rectangular, poorly setose anteriorly. Article 5 of
pereopods 1-2 setose posteroproximally; article 2
of pereopod 3 of broad form but tapering distally,
articles 4?5 of pereopods 3-4 narrow to medium,
article 2 of pereopods i-A setose posteriorly;
pereopod 5 ordinary, article 2 strongly setose ven-
trally, article 3 ordinary, dactyl normal. Epimera
1-2 bearing numerous long posterior setae, without
midfacial setae above ventral facial ridge; epimeron
3 ordinary. Urosomite 1 generally naked except for
sparse apicoventral setae or spines near base of uro-
pod 1; urosomite 3 without dorsal hook or special
process. Peduncle of uropod 1 normally elongate,
without apicoventral spike, with special enlarged
apicomedial spine; peduncular apices of uropods 1-2
not combed; inner ramus of uropod 1 with mar-
ginal spines in one row, no rami continuously
spinose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, article 2 of outer ramus carry-
ing 2-4 short to medium apical setae. Telson with
2 or more apical spines or setae on each lobe plus
setules, with special dorsal and lateral spines or
setae.
DESCRIPTION.?Rostrum fully developed, uncon-
stricted to constricted. Fuzz on article 1 of antenna
1 in male present; calceoli on male primary flagel-
lum of antenna 1 absent as far as known. Calceoli
of male antenna 2 absent as far as known. Prebuc-
cal parts ordinary, poorly separated from each
other, upper lip dominant. Right lacinia mobilis
bifid, thin; mandibular palp thin, article 1 short,
article 2 without outer setae, apex of article 3
oblique. Lower lip bearing cones. Outer plate of
maxilla 1 with 11 spines, one spine especially
thickened. Inner plates of maxilliped thick, ordi-
narily setose. Coxae 2-4 with weak anterodorsal
humps. All posterior spines on article 6 of pereo-
pods 1-2 thick and stiff, midapical spine present.
Article 2 of pereopod 5 without facial setae, bearing
only one ridge. Peduncle of uropod 1 with dorso-
lateral spines confined apically, medial spines
widely spread; peduncle of uropod 2 with only one
medial setule confined apically. Peduncle of uropod
3 lacking extra subapical setae or spines. Telson
with ordinary pair of midlateral or dorsal setules
on each side.
TYPE-SPECIES.?Phoxus villosus Haswell, 1879
[original designation].
COMPOSITION.?Parharpinia warte, new species.
RELATIONSHIP.?Parharpinia bears the special
apicomedial spine on the peduncle of uropod 1
and the shortened article 2 of antenna 1 as in
Pontharpinia but in almost all other characters (ex-
cept telson, epimera 1-2 and article 2 of pereopod
3) it resembles Birubius. It differs from Birubius
in the special spine on uropod 1, the short article
2 of antenna 1 with especially widespread setation,
the posteriorly setose epimera 1-2, the tapering
article 2 of pereopod 3, the posteriorly setose article
2 of pereopods 3-4, and the presence of supernu-
merary dorsal and apical spination and setation on
the telson. The facial spines on article 4 of antenna
2 are mostly condensed into one main row in con-
trast to Pontharpinia and Birubius. The maxilli-
pedal nail is poorly developed in contrast to Biru-
bius.
Parharpinia resembles Mandibulophoxus and
Basuto in the special spine of uropod 1 and the
short article 2 of antenna 1 but differs from those
genera in the relatively small palpar hump on the
mandible, the better developed molar, the small
gnathopod 2, the well setose article 2 of pereopods
3-4, the large article 5 of antenna 2, the dorsal
176 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
telsonic spination, the posteriorly setose epimera
1-2, and the poor nail on the maxilliped.
Parharpinia resembles Leongathus in article 2
of pereopod 3, epimeral setation, facial spination
on article 4 of antenna 2, the weak nail of the max-
illiped, the slightly ensiform antenna 2, and the
slightly enlarged articles 4-5 of pereopod 5 but
Parharpinia bears a special medial spine on the
peduncle of uropod 1, has a small gnathopod 2,
and a reduced and spinose molar.
If any evolutionary flow occurred between Par-
harpinia and Birubitts it might be marked in B.
karobrani and B. babaneekus through the ventrally
setose article 2 of pereopod 5 or in B. elecbanus
through the truncate telsonic lobes and cuticle.
Those species of Birubius, however, remain fully
distinguished by the several good generic characters
stated above.
Key to the Species of Parharpinia
Bulbar setules of cuticle set in space void of villi P. watte
Bulbar setules and villi of cuticle evenly scattered, no voids P. vttlota
Parharpinia villosa (Haswell)
FIGURES 72-78
Phoxus villosus Haswell. 1879:258-259, pi. 9: fig. 2: 1882:
236-237.
Parharpinia villosa.?Stebbing, 1906:147-148 [in part); 1910:
635.
Paraphoxus villosus.?J. L. Barnard, I960: 285-285.
Not Parharpinia villosa.?Tattersall, 1922:4-6, pi. 1: figs. 7-14
[- Brolgus tattcrrs<i//i].-Schellenberg, 1926:300-301 [-
Parharpinia? sp.]: 1931:75-78, fig. S9 [= P. sinuatus].
Not Pontharpinia villosa.?K. H. Barnard, 1940:442-443
[?? Pontharpinia? sp].
Not Phoxus batei, Thomson, 1882:232-233, pi. 17: fig. 2 [not
Haswell, 1880a] [= Protophoxus australis].
DESCRIPTION OF FEMALE.?Head about 20 percent
of total body length, greatest width about 75 per-
cent of length; rostrum unconstricted, broad, reach-
ing middle of article 2 on antenna 1. Eyes small
to medium, stained pink, clear of occluding pig-
ment, ommatidia especially small. Article 1 on pe-
duncle of antenna 1 about 1.8 times as long as
wide, about twice as wide as article 2, ventral mar-
gin with 2-3 setules, produced dorsal apex with 5
setules; article 2 about half as long as article 1,
with two ventral rows of 17 thick and thin setae;
primary flagellum with 13-15 articles, about 0.8
times as long as peduncle, bearing aesthetascs; ac-
cessory flagellum with 11-13 articles. Spine for-
mula on article 4 of antenna 2 = 1-3-8 or 1-3-10
(or other variations), dorsal margin with notch
bearing 2-3 setae and 1-2 spines, ventral margin
densely setose, with one main ventrodistal short
spine and up to 4 other thinner spines; article 5
about 0.95 times as long as article 4, facial spine
formula = 2-2-2-2-1 or 2-2-2-2-2-2 or 11-12 in
tandem alternating long and short (or other varia-
tions), dorsal margin naked, ventral margin with
8 sets of 1-2 long to short setae and spines, 4 ven-
trodistal long to short spines, 2 of these placed as
subdistal facial spines; flagellum about 0.85 times
as long as articles 4-5 of peduncle combined, with
13-15 articles. Mandibles with weak to medium
palpar hump; right incisor with 3 teeth and faint
notches; left incisor with 3 humps in 2 branches;
right lacinia mobilis bifid, distal branch shorter
than proximal, distal branch broad, subbifid, den-
ticulate, proximal branch simple, pointed; left la-
cinia mobilis with 5 teeth; right rakers 8-11; left
rakers 9-11; molar in form of elongate bulbous
hump demarcated mainly by spines, each molar
with 7-9 primarily medium spines plus one short
spine strongly disjunct; palp article 1 short, article
2 with 3 long inner apical setae and 3-8 other long
and short inner setae, article 3 about 1.1 times as
long as article 2, oblique apex with 12-14 spine-setae,
basofacial formula = 0-1 or 1-1. Inner plate of
maxilla 1 large, broad, bearing one long apical
pluseta, one similar apicomedial seta, 2 apicolateral
much shorter setae; palp article 2 with 8-10 apical-
medial marginal spines and submarginal setae. Inner
plate of maxilla 2 shorter and narrower than outer,
outer with 3-6 apicolateral setae, inner with 2 medial
setae. Inner plate of maxilliped with 2-6 large thin
apical spines, 4 apicofacial setae, 2 medial setae;
outer plate with 8-12 medial and apical spines, 9-13
apicolateral setae; palp article 1 lacking apicolat-
eral seta, article 2 with 2 apicolateral setae, medial
margin of article 2 strongly setose, article 3 weakly
NUMBER 245 177
FIGURE 72.?Parharpinia villosa (Haswell), female "a," 16.3 mm (b =? male "b," 9.5 mm; / ?
female "f," 11.9 mm; n ? lectotype, female "n," 14.8 mm; u = juvenile "u," 8.8 mm; y ? male
"y," 10.3 mm).
178 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
P2
FIGURE 73.?Parharpinia villosa (Haswell), female "a," I6.S mm (b - male "b," 9.5 mm; / ?
female "f." 11.9 mm; / - male "j ," 9.8 mm; n - lectotype, female "n," 14.8 mm).
NUMBER 245 179
W
FIGURE 74.?Parharpinia villosa (Haswell), female "a," 16.3 mm (6 ? male "b," 9.5 mm; / <
female "t," 11.9 mm; n ? lecototype, female "n," 14.8 mm).
protuberant, with 11 scattered facial setae, 2 groups
of 1-4 lateral setae, nail of article 4 short, with one
accessory setule. Coxa 1 expanded distally, anterior
margin concave; main ventral setae of coxae 1-4 =
20-11-14-35, posteriormost seta of coxae 1-3 short-
ened; anterior and posterior margins of coxa 4 di-
vergent, posterior margin oblique, almost straight,
setose, posterodorsal corner sharp-rounded, postero-
dorsal margin short, concave, width-length ratio
of coxa 4 = 8:7. Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 = 2-6-5-5,
long anteriors = 4-7-0-0, short anteriors = 4?5-2-4,
no others. Gnathopods ordinary; width ratios of
articles 5-6 on gnathopods 1-2 = 28:36 and 29:37,
length ratios = 62:67 and 56:62; palmar humps
ordinary, palms oblique; article 5 of gnathopod 1
weakly elongate, ovate, posterior margin flat; arti-
cle 5 of gnathopod 2 trapezoidal, posterior margin
rounded, almost lobate. Pereopods 1-2 similar, fa-
cial setae formula on article 4 = (5-8), on article
180 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
bR3
FIGURE 75.?Parharpinia villosa (Haswell), upper: juvenile "m," 3.9 mm; lower: female "a," 16.S
mm (b = male "b," 9.5 mm).
NUMBER 245 181
Al
FIGURE 76.?Parharpinia villosa (Haswell), juvenile "m," S.9 mm.
5 = (5-7); main spine of article 5 extending to M.
90 on article 6, article 5 lacking proximoposterior
spines; spine formula of article 6 = 6 + 6 or 7 + 7
plus middistal seta or spinule; acclivity on inner
margin of dactyls of pereopods 1?2 weak to obso-
lescent, emergent setule long to short, midfacial
pluseta very short. Coxae 5-7 posteroventral seta
formula = 21-13-14. Articles 4-5 of pereopods 3-4
narrow, facial spine rows dense, facial ridge for-
mula on article 2 of pereopods 3-5 = 0-1-1, article
2 of pereopod 3 tapering distally; width ratios of
articles 2, 4, 5, 6 of pereopod 3=41:25:24:13,
of pereopod 4 = 60:26:20:10, of pereopod 5 =
80:20:15:8, length ratios of pereopod 3 = 67:31:
35:36, of pereopod 4 = 76:50:53:66, of pereopod
5 = 84:22:21:27; article 2 of pereopod 5 reach-
ing middle of article 4, posterior and ventral
margins of article 2 on pereopods 3-5 setose, medial
apex of article 6 finely combed, bearing 4 vestigial
digital processes. Posteroventral corner of epimeron
182 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 77.?Parharpinia villosa (Haswell), juvenile "h," S.7 mm (m - juvenile "m," S.9 mm).
NUMBER 245 183
P5
FIGURE 78.?Parharpinia viliosa (Haswell), juvenile "h," S.7 mm.
1 rounded, posterior margin weakly convex, ser-
rate, setose, anterior and ventral margins with 8-10
long to medium setae; posteroventral corner of epi-
meron 2 rounded, posterior margin almost straight,
serrate, setose, facial setae = 9-11, some middle
pairs set vertically or irregularly, somewhat
crowded forward; posteroventral corner of epime-
ron 3 rounded, posterior margin weakly convex,
serrate, setose, ventral margin naked, face with 3-5
scattered spines and 1-3 setules near ventral mar-
gin in middle. Urosomite 1 with 1-3 ventral spines
at base of uropod 1, one midventral seta, articula-
tion line complete; urosomite 3 weakly protuberant
dorsally. Rami of uropods 1-2 with articulate en-
larged apical nails, outer rami of uropods 1-2 with
gins naked or with 1-2 setae, article 2 of outer ramus
of uropod 1 with 5-10 dorsal spines, inner with 2,
outer ramus of uropod 2 with 4-7 dorsal spines,
inner with none; peduncle of uropod 1 with 3
apicolateral spines and 3-5 basofacial spinules, me-
dially with 7-9 marginal spines, apicalmost spine
set inward on end of peduncle medially, unenlarged;
peduncle of uropod 2 with 8-10 dorsal spines,
medially with one small apical setule. Peduncle
of uropod 3 with 6-8 ventral spines, dorsally with
2 lateral spines, 1-2 medial spines; rami feminine,
inner extending to M. 75 on article 1 of outer ramus,
apex with 2-6 setae, apicomedial and lateral mar-
gins naked or with 1-2 setae, article 2 of outer ramus
short, 0.10, bearing 2-4 short to medium setae,
medial margin with 2-6 setae, apicomedial margin
of article 1 with 3 setae, lateral margin with 4-6
acclivities, spine formula = 2-2-2-2(-2-2) (paired
long and short), setal formula = 0 or 0-0-0-0-0-0-3.
184 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Telson ordinary, length-width ratio =1:1 or 13:
12, not fully cleft, each apex wide, rounded to trun-
cate, lateral acclivity broad, shallow, with 1-2 lateral
and 1-2 medial spines separated by short setule,
spines variable, short to long, dorsodistal surface
with 2-4 additional setal spines on each lobe, mid-
lateral setules diverse. Cuticle closely shagreened,
villi grouped as ripples, setules sparse.
DESCRIPTION OF MALE.?Eyes often distending
head; rostrum shorter than in female. Article 2
of antenna 1 with fewer ventral setae than in fe-
male. Facial spine formula on article 4 of antenna
2 = 1-2-8 (= 1-2-3-5) or 1-2-9 or 1-2-10, on
article 5 = 2-2-2-3 or 2-2-2 or 2-1-2-2 or 2-2-2-2;
article 5 with 3-4 dorsal sets of male setae (hidden
in illustration), no calceoli, ventrodistal apex with
2 thin spines, dorsodistal pair of female absent;
flagellum with 27-30 articles, no calceoli. Baso-
facial setal formula on article 3 of mandibular
palp = 1-2 + 1-3 midfacial setae, terminal = 10-12;
distal branch of right lacinia mobilis as in female
or broader and more deeply bifid. Article 2 of only
pereopod 5 narrower than in female. Epimera 1-3
broadened, posterior margin of epimeron 3 short-
ened; setal formulas, epimeron 1 ventral = 10, epi-
meron 2 facial = 7, epimeron 3 posterior = 9, fa-
cial = 3. Spine formulas of uropods, uropod 1 pe-
duncle apicolateral = 3, basofacial = 3-4, medial =
6, uropod 2 peduncle dorsal = 8, dorsal spines on
outer ramus of uropod 1 = 5, of uropod 2 = 4-5,
inner ramus of uropod 1 = 2 , of uropod 2 = 0.
Spine formula on uropod 3, article 1 of outer
ramus = 0-0-1-1-1-1, setal formula = 1?1?1?1?1?1.
Telson elongate, distal spines shortened and fewer.
OBSERVATIONS.?Spine formula on face of article
4 on antenna 2 subject to interpretation, generally
quoted here as bearing 3 rows, first row with one
apicomedial spine, next row of 2 on face and next
row representing agglomeration of 2 weakly dis-
tinct rows, sometimes divided weakly into subequal
groups, sometimes merging imperceptibly, some-
times with skewed members; one subadult female
from WPBES T46-62 with single, heavily thick-
ened midfacial spine on epimeron 3.
DESCRIPTION OF JUVENILE.?Juvenile "h," 3.7
mm: Highly distinctive. Eyes small. Article 2 of
antenna 1 with one long apicoventral spine, pri-
mary flagellum with 4 articles, accessory flagellum
with 2. Article 4 of antenna 2 with one long ven-
tral spine-seta, facial formula = 1-2; article 5 with
one dorsal spinule, 2 ventral setules, one long api-
coventral seta and one spine; flagellum with 4 arti-
cles. Right lacinia mobilis bifid, with 2 equally ex-
tending branches, distal slightly broadened; left
lacinia mobilis with 4 teeth, middle pair shortened;
rakers 6 on both sides; molars with 3-4 spines plus
one short disjunct spine; palp article 2 naked, arti-
cle 3 with 3 apical spine-setae, otherwise naked.
Outer plate of maxilla 1 with only 6 spines; palp
with 2 apical setae. Setation reduced on maxilli-
peds, gnathopods, pereopods and coxae. Setal for-
mula of coxae 1-4 - 1-1-1-0 (latter with 2 setules
as usual). Articles 4-5 of pereopods 1-2 lacking
facial and posterior setae except for one apicopos-
terior seta on article 4; article 5 with 2 distal
spines; article 6 with rows of one and one spine.
Pereopods 3-5 simplified, article 2 with only one
seta each, article 2 of pereopod 3 only weakly ta-
pering. Epimera 1-3 with few large posterior ser-
rations, one large posterodoi sal seta each, no other
setae. Peduncle of uropod I lacking basofacial
setae, with one apicolateral spine, uropod 2 similar,
outer rami of uropods 1-2 each with one accessory
apical nail, otherwise all rami naked except for
ordinary apical nails. Uropod 3 with shorter inner
ramus and longer article 2 on outer ramus than in
adult, setae on article 1 of outer ramus reduced to
one, dorsal margins of uropods with pectinations.
Telson short, each lobe with narrow apex bearing
2 setules.
Juvenile "m," 3.9 mm: More mature than 3.7
mm juvenile "h," setae and spines more numerous,
apical nails on uropods 1-2 absent, these spines
apparently migrant to dorsal margins (see illustra-
tions).
Juvenile "k," 3.0 mm: Uropods 1-2 like those of
3.7 mm juvenile "h".
Juvenile "p," 1.7 mm: Uropods 1-2 lacking any
but apical nails, no other spines.
ILLUSTRATIONS.?Outline drawings of gnathopods
from lectotype and females "a" and "f given to
show variations. Inner plate of maxilliped, uropod
3 and telson of female "f" more highly magnified
than views from other specimens. Juvenile "h,"
3.7 mm: head damaged, illustration based on par-
tial restoration; left mandible missing; nail of max-
illipedal palp article 4 scarcely distinct. Juvenile
"m," 3.9 mm: following parts as in 3.7 mm juve-
nile "h": upper and lower lips, maxilla 2, dactyls
of pereopods 1-5, telson, cuticle and maxilliped,
NUMBER 245 185
but latter with one lateral seta on outer plate, palp
articles 2-3 with several additional setae; molars
of mandibles drawn inverted, each body of man-
dible with ordinary protrusion at base of palp;
pereopod 4 illustrated in 2 pieces from right and
left members; views of head and pereopods 3-5
partially reconstructed; illustration of urosome
slanting away from observer; shapes of coxae 1?4
similar to those of 3.7 mm juvenile "h" but setal
formula = 5-4-3-2.
LECTOTYPE.?AM, female "n," 14.8 mm, parts on
5 slides (this specimen, apparently labeled "Type"
long after being described by Haswell (see J. L.
Barnard, 1974), was presumably dissected by K.
Sheard and the various appendages mounted by
E. F. Gurjanova; all appendages have now been
remounted by us and returned to AM).
TYPE-LOCALITY.?AM G. 5413, Port Jackson, New
South Wales, Australia.
VOUCHER MATERIAL (all illustrated).?CPBS
22N/2, female "a," 16.3 mm; CPBS 52N/1171,
male "b." 9.5 mm; CPBS 51S/4, female "?," 11.9
mm; RHM, male " j , " 9.8 mm; CPBS41N/4, juve-
nile "u," 8.8 mm; AM P. 18312, Hood Point, SW
Australia, 145 m, male "y," 10.3 mm; CPBS 26S/2,
juvenile "h," 3.7 mm; PPBES 985/4: juvenile "k,"
3.0 mm; juvenile "m," 3.9 mm, juvenile "p," 1.7
mm.
REMARKS.?The lectotype, an adult female, can
no longer be examined in toto, but consists of a
series of permanent mounts of the dissected ap-
pendages and mouthparts. It has been used exten-
sively in making the fuller description of this spe-
cies. Upper and lower lips are missing; some ap-
pendages are incomplete; many spines and setae
are broken or missing altogether. The only brood
lamella, remaining more or less intact, is setose.
Specimen "a," a female not fully adult, from which
the bulk of the illustrations has been drawn, differs
essentially from female "f" and the lectotype in
quantitative detail only: the number of individual
setae, spines and of spine-seta clusters is fewer on
some appendages; spines on article 5 of antenna 2
are shorter and on the inner plate of the maxil-
liped are fewer than in the adult; uropod 3 lacks
the full complement of setae on both rami; adult
appendages differing in any but most minor details
from those of female "a" have been separately
illustrated.
Detailed comparison of the size of lectotype ap-
pendages with those of females "a" and "f" (articles
5 and 6 of gnathopod 1 illustrated in outline as
an example) indicate that Haswell's specimen must
have been larger than female "a." Using the tech-
nique adopted in this investigation for measuring
length (viz., along a line midway between dorsal
and ventral, from tip of rostrum to base of telson),
the lectotype would be more than 16.3 mm long,
considerably in excess of the 7 lines estimated by
Haswell.
MATERIAL.?AM, Lectotype, Port Jackson (1),
Hood Point (1); CPBS, 21 samples from 10 stations
(29); WPBES, 2 samples from 2 stations (2); RHM,
2 samples from 2 stations (2); PPBES, 2 samples
from one station (2); SBS, one sample, dredged (1);
Macreadie, 2 samples, plankton (2).
DISTRIBUTION.?New South Wales, Sydney, to Vic-
toria, Port Phillip Bay; Western Australia, Hood
Point; depth range, 8-145 m; sediments: sand,
shelly sand, sand and gravel, sand and mud.
Parharpinia watte, new species
FIGURES 79-81
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total body length, greatest width about 65
percent of length; rostrum scarcely constricted,
somewhat short, reaching apex of article 1 on an-
tenna 1. Eyes small, stained pale eosin, clear of
occluding pigment, ommatidia small. Article 1 on
peduncle of antenna 1 about 1.7 times as long as
wide, about twice as wide as article 2, ventral mar-
gin with about 5 setules, weakly produced dorsal
apex with one setule; article 2 about 0.5 times as
long as article 1, with apicoventral cycles of 10-11
setae; primary flagellum with 11 articles, about as
long as peduncle, lacking aesthetascs, [accessory
flagellum unknown, broken]. Spine formula on
article 4 of antenna 2 = 1-3-4-5 (or 1-3-9), dorsal
margin with notch bearing 2 setae and one spine,
ventral margin with 3 groups of 1-2 long to me-
dium setae, one ventrodistal long spine; article 5
about 0.85 times as long as article 4, facial spine
formula = 2-2-2-2 or 2-2-2-2-1, dorsal margin
naked, ventral margin with 3-4 sets of 1-3 long to
medium setae, 2 ventrodistal medium spines, one
subdistal facial spine counted in facial formula;
flagellum about 1.1 times as long as articles 4-5
of peduncle combined, with 10 articles. Mandibles
186
SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY
Al I , A2
FIGURE 79.?Parharpinia warte, new species, holotype, female "a," 9.6 mm.
with weak pal par hump; right incisor with 3 teeth; narrow, proximal branch simple, pointed, with
left incisor with 3 humps in 2 branches; right marginal denticles and facial hump forming cusp
lacinia mobilis bifid, distal branch much shorter in crotch; left lacinia mobilis with 4 teeth plus one
than proximal, distal branch erect and thorn-like, accessory tooth, middle teeth lengthened; right and
NUMBER X45 187
P5
FIGURE 80.?Parharpinia warte, new species, holotype, female "a," 9.6 mm.
left rakers 9; molar in form of elongate bulbous
hump demarcated mainly by spines, right molar
with 10 primarily medium spines plus 1-2 rudi-
mentaries, plus one spine strongly disjunct, left
molar with 9 primarily medium spines plus one
spine strongly disjunct, plume absent; palp article
1 short, article 2 with one medium inner apical seta
and 2 other shorter inner setae, article 3 about as
long as article 2, oblique apex with 8 spine-setae,
basofacial formula = 0-1. Inner plate of maxilla 1
large, broad, bearing or lacking one long apical
pluseta, one shorter apicomedial seta, 3 apicolateral
188 SMITHSONIAN CONTRIBUTIONS TO ZOOLOG*
FIGURE 81.?Parharpinia warte, new species, holotype, female "a," 9.6 mm.
much shorter setae; palp article 2 with 6 apical-
medial marginal setae. Plates of maxilla 2
extending subequally, outer broader than inner,
outer with 4 apicolateral setae, inner with 3 medial
setae. Inner plate of maxilliped with 2 large thin
apical spines, 4 apicofacial setae, 2 medial setae;
outer plate with 8 medial and apical spines, 5 apico-
lateral setae; palp article 1 lacking apicolateral seta,
article 2 with one apicolateral seta, medial margin
of article 2 moderately setose, article 3 with 6 facial
setae, one lateral seta, article 4 with 2 widely spaced
apical accessory setules, nail absent. Coxa 1 strongly
expanded distally, anterior margin weakly concave;
main ventral setae of coxae 1-4 = 11?7?6?6, poste-
riormost seta of coxae 1-4 shortest; anterior and
posterior margins of coxa 4 divergent, posterior
margin oblique, straight, posterodorsal corner
sharp-rounded, posterodorsal margin short, con-
cave, width-length ratio of coxa 4 = 7:8. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 2-4-(3^)-(3-4), long anteriors =
6-6-0-0, short anteriors = 3-3-3-2, no others. Gna-
thopods ordinary; width ratios of articles 5-6 on
gnathopods 1-2 = 24:38 and 27:40, length ratios =
62:62 and 52:64; palmar humps ordinary, palms
oblique; article 5 of gnathopod 1 ovate, posterior
margin rounded-flat, short; article 5 of gnathopod
2 subtriangular, posterior margin rounded, short,
NUMBER 245 189
almost lobate. Pereopods 1-2 similar; facial setae
formula on article 4 = 2 and 2, on article 5 = 3 and
3; main spine of article 5 extending to M. 85 on
article 6, article 5 lacking proximoposterior spines;
spine formula of article 6 = 4 + 6 plus middistal
seta, one spine especially long; acclivity on inner
margin of dactyls of pereopods 1-2 weak, emergent
setule short, midfacial pluseta ordinary. Coxae 5-7
posteroventral setule formula = 6-5-6. Articles 4-5
of pereopods 3-4 narrow, facial spine rows sparse,
facial ridge formula on article 2 of pereopods 3-5 =
0-1-1, (pereopod 5 with additional short distomar-
ginal ridge); width ratios of articles 2, 4, 5, 6 of
pereopod 3 = 49:30:28:16, of pereopod 4 = 70:32:
25:13, of pereopod 5 = 95:22:20:10, length ratios of
pereopod 3 = 81:37:39:47, of pereopod 4 = 93:67:
68:88, of pereopod 5 = 110:23:27:36; article 2 of
pereopod 5 exceeding apex of article 4, ventral mar-
gin setose, posterior margins of article 2 on pereo-
pods 3-4 setose, article 2 of pereopod 3 tapering
distally; medial apex on article 6 of pereopod 5
finely combed, bearing 3 short digital processes.
Posteroventral corner of epimeron 1 rounded, pos-
terior margin straight, serrate, setose, anteroventral
margin with 3 medium setae, posteroventral mar-
gin with 2 medium setae; posteroventral corner of
epimeron 2 rounded to weakly protuberant, with
small, sharp tooth, posterior margin straight, ser-
rate, setose, facial setae = 5 crowded anteriorly;
posteroventral corner of epimeron 3 weakly protu-
berant, with sinus, with small tooth, posterior mar-
gin straight, weakly serrate, setose, face with hori-
zontal row of 2 short spines; pleonites 1-3 each
with 3 dorsal longitudinal rugosities. Urosomite 1
with ventral spine at base of uropod 1, articulation
line almost complete; urosomite 3 weakly protuber-
ant dorsally. Rami of uropods 1-2 with articulate
enlarged apical nails, outer ramus of uropod 1 with
4 dorsal spines, inner with 2, outer ramus of uropod
2 with 4 dorsal spines, inner lacking dorsomedial
spines; peduncle of uropod 1 with 3 apicolateral
spines and 4 short dispersed basofacial setae, me-
dially with 5 marginal spines, apicalmost enlarged
but set inward; peduncle of uropod 2 with 7 dorsal
spines, medially with one tiny apical spine. Pedun-
cle of uropod 3 with 4 ventral spines, dorsally with
one lateral spine, one medial spine and setule; rami
feminine, inner extending to M. 70 on article 1 of
outer ramus, apex with one seta, medial and lateral
margins naked, article 2 of outer ramus ordinary,
0.20, bearing 2 medium setae, medial apex of article
1 with 2 setae, lateral margin with 3 acclivities,
spine formula = 1-1-2-1, setal formula = 0-0-0-1.
Telson ordinary, length-width ratio = 11:13, not
fully cleft, each apex wide, truncate, lateral accliv-
ity absent, with lateral short and medial long, thin
spine separated by long setule, each lobe dorsally
with spinule at M. 70, midlateral setules large,
diverse. Cuticle with bulbar setules at edges of clear
spaces in midst of dense pebble knobs, dorsally
organized on pleon into alternate rows of blunt
and sharp knobs, emergent setules especially long,
branched.
OBSERVATIONS.?Head damaged but also asym-
metrically formed, left side deeper than right; pos-
terior half of sternite 6 heavily calcified and form-
ing half-ring below body; article 3 of antenna 2
with apicodorsal spine.
ILLUSTRATIONS?Some rakers of mandibles omit-
ted from drawings.
HOLOTYPE.?AM, female "a," 9.6 mm. Unique.
TYPE-LOCALITY.?AM P. 18276, 5 Jul 1962, Great
Australian Bight, south of Eucla, Western Austra-
lia, 33?05' S, 128?40' E, 83 m, bottom unknown.
RELATIONSHIP.?This species is very closely simi-
lar to Parharpinia villosa and can be readily dis-
tinguished only by the cuticular structures. Par-
harpinia villosa has evenly distributed villi and
bulbar setules lacking any marked clear space
around them, whereas P. warte has tightly packed
villi, apically blunted and forming a ring around
spaces in which lie the bulbar setules. The unique
holotype of P. warte might be an aberrant specimen
of P. villosa in all other respects. Various minor
distinctions in setal densities and sizes of gnatho-
podal articles might be explained on the aberrancy
theory but the odd cuticle cannot be so explained.
MATERIAL.?AM, one sample (1).
DISTRIBUTION.?Great Australian Bight, Eucla.
83 m.
Protophoxus K. H. Barnard
Protophoxus K. H. Barnard, 1930:335.?Barnard and Drum-
mond, 1976:532.
DIAGNOSIS OF MALE.?Eyes present. Flagella of
antennae 1-2 unreduced in male. Article 2 of an-
tenna 1 especially shortened, ventral setae widely
spread. Article 1 of antenna 2 not ensiform, article
190 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
3 with 2 setules, facial spines on article 4 primarily
in one main row, article 5 ordinary in size. Right
mandibular incisor with 3 teeth; molar not tritura-
tive, small, pillow-shaped, bearing 4 or more
splayed, semiarticulate spines, usually bearing fuzz;
palpar hump medium. Palp of maxilla 1 biarticu-
late; inner plate with 4 setae. Setation of maxilla 2
ordinary. Inner plate of maxillipeds ordinary; apex
of palp article 3 not or weakly protuberant, dactyl
elongate, apical nail distinct, medium. Gnathopods
ordinary, small, similar; article 5 of gnathopods 1-2
of ordinary length, free, without eusirid attach-
ment; palms oblique, hands of gnathopods 1-2
ordinary, ovatorectangular, poorly setose anteriorly.
Article 5 of pereopods 1-2 bearing posteroproximal
setae. Article 2 of pereopod 3 of broad form but
tapering distally, articles 4-5 of pereopods 3-4
broad to medium, article 2 of pereopods 3-4 not
setose posteriorly; pereopod 5 ordinary, article 2
naked ventrally, article 3 ordinary; dactyl normal.
Epimera 1-2 lacking long posterior setae, without
midfacial setae above ventral facial ridge; epimeron
3 ordinary. Urosomite 1 generally naked; urosomite
3 without dorsal hook or special process. Peduncle
of uropod 1 normally elongate, without apicoven-
tral spike, with special enlarged apicolateral spine,
only peduncular apices of uropod 2 combed; inner
ramus of uropod 1 with one row of marginal spines,
no rami continuously spinose to apex, inner ramus
of uropod 2 ordinary. Uropod 3 highly elongate,
article 2 of outer ramus carrying 2 long apical
setae. Telson with 2 or more apical spines, with
special dorsal spines.
DESCRIPTION.?Rostrum fully developed. Fuzz on
article 1 of antenna 1 in male present; calceoli on
male primary flagellum of antenna 1 present. Cal-
ceoli on article 5 of male antenna 2 present; flagel-
lum in male with calceoli. Prebuccal parts ordinary,
poorly separated from each other, upper lip domi-
nant. Right lacinia mobilis bifid, thin; mandibular
palp medium to thin, article 1 short, article 2 with-
out outer setae, apex of article 3 oblique. Lower
lip bearing cones. Outer plate of maxilla 1 with 11
spines, one spine especially thickened. Inner plates
of maxilliped thick, ordinarily setose. Coxae 2-4
without special antero-dorsal humps. All posterior
spines on article 6 of pereopods 1-2 thick and stiff,
midapical spine present. Article 2 of pereopod 5
without facial setae. Peduncle of uropod 1 with
dorsolateral spines widely spread, medial spines
widely spread; peduncle of uropod 2 with only one
medial spine confined apically. Peduncle of uropod
3 lacking extra subapical setae or spines. Telson
with ordinary pair of midlateral or dorsal setules
on each side.
TYPE-SPECIES.?Protophoxus australis K. H.
Barnard, 1930 (monotypy).
COMPOSITION.?Unique.
RELATIONSHIP.?Protophoxus shares with Par-
harpinia the supernumerary dorsal spination on the
telson, the presence of a special spine on the pedun-
cle of uropod 1, and the strong taper of article 2 on
pereopod 3. Protophoxus differs from Parharpinia
in the shift of the special spine on the peduncle of
uropod 1 to the lateral side, the absence of posterior
setation on epimera 1-2, the absence of strong ven-
tral setation on article 2 of pereopod 5, and the
more strongly distinct nail of the maxillipedal
dactyl.
Protophoxus and Parharpinia share thin articles
4?5 of pereopods 3-4 but this character is almost
as well developed in some of the thin membered
species of Birubius.
Protophoxus australis K. H. Barnard
Phoxus batei.?Thomson, 1882:232-233, pi. 17: fig. 2 [not
Haswell).
Protophoxus australis K. H. Barnard, 1930:335-336, fig. 12.
?Barnard and Drummond, 1976:534, fig. 4 [part].
Pontharpinia australis.?Hurley, 1954:581-587. figs. 1-28.
DISTRIBUTION.?New Zealand, from Three Kings
Islands south to Stewart Island, neritic, 0-3 m, con-
firmed benthic 13-37 m.
BIRUBIINAE
DIAGNOSIS.?Article 2 of antenna 1 ordinary to
elongate; mandibular molar reduced to a small
hump with articulate spines; palp of maxilla 1
biarticulate; setation on maxilla 2 ordinary; gna-
thopod 2 as small as gnathopod 1; article 2 of pereo-
pod 3 of broad form, not tapering distally; pereo-
pod 5 ordinary.
DESCRIPTION.?Article 5 of antenna 2 of normal
size; epimeron 3 of nonrounded classification; apices
of peduncles on uropods 1-2 not combed.
TYPE GENUS.?Birubius, new genus. (Birubius is
NUMBER 245 191
picked as type genus of the subfamily because we
are not certain that Metharpinia, the oldest name,
belongs to the subfamily.)
COMPOSITION.?Kulgaphoxus, new genus; Micro-
phoxus J. L. Barnard; Tickalerus, new genus; Yan,
new genus; possibly Metharpinia Schellenberg.
Key to the Genera of Birubiinae
1. Urosomite S with large dorsal hook on each side, outer ramus of uropod 3 severely reduced
in size, not longer than peduncle 2
Urosomite S lacking dorsal hooks, outer ramus of uropod S usually more than 1.5 times as
long as peduncle S
2. Uropods 1-2 lacking accessory apical nails on inner rami, ventral setae on article 2 of antenna
1 placed in middle Tickalerus, new genus
Uropodt 1-2 bearing accessory apical nails on inner rami, ventral setae on article 2 of
antenna 1 situated proximally Kulgaphoxus, new genus
3. Dactyl of pereopod 5 vestigial Yan, new genus
Dactyl of pereopod 5 fully formed 4
4. Telson lacking supernumerary lateral spines and setae except for normal pair of midlateral
and dorsal sctules on each side 5
Telson with supernumerary lateral spines and setae besides normal pair of midlateral and
dorsal setules on each side 6
5. Gnathopod 2 significantly larger than gnathopod 1 . . . Cunmurra, new genus (see Brolginae)
Gnathopods 1-2 subequal in size Birubhu
6. Coxa 4 with posterior setae, see text Metharpinia
Coxa 4 lacking posterior setae, see text Microphoxus
Birubhu Barnard and Drummond
Birubius Barnard and Drummond, 1976:543.
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
ordinary to elongate, ventral setae widely spread.
Article 1 of antenna 2 not or weakly ensiform; arti-
cle 3 with 2 setules; facial spines on article 4 in 2
or more rows; article 5 ordinary in size. Right man-
dibular incisor with 3-4 teeth; molar not triturative,
ordinary, pillow-shaped, bearing 4 or more splayed
semiarticulate spines, usually bearing fuzz; palpar
hump small to medium. Palp of maxilla 1 biarticu-
late, inner plate with 3-4 setae. Setation of maxilla
2 ordinary. Inner plate of maxillipeds ordinary;
apex of palp article 3 not or weakly protuberant,
dactyl elongate, apical nail distinct, medium to
elongate. Gnathopods ordinary, small, similar; arti-
cle 5 of gnathopods 1-2 of ordinary length, free on
gnathopod 2, often elongate on gnathopod 1, with-
out eusirid attachment; palms oblique, hands of
gnathopods 1-2 ordinary, ovatorectangular, poorly
setose anteriorly. Article 5 of pereopods 1-2 bear-
ing setae posteroproximally. Article 3 of pereopod 3
of broad form, rarely and scarcely tapering distally,
articles 4-5 of pereopods 3-4 broad to narrow, arti-
cle 2 of pereopods 3-4 not setose posteriorly; pereo-
pod 5 ordinary, article 2 naked or strongly setose
and toothed ventrally, article 3 ordinary, dactyl
normal. Epimera 1-2 lacking numerous long pos-
terior setae, without midfacial setae above ventral
facial ridge; epimeron 3 ordinary. Urosomite 1 gen-
erally naked except for sparse apicoventral setae or
spines near base of uropod 1, rarely with lateral
line of setae; urosomite 3 without dorsal hook or
special process. Peduncle of uropod 1 normally
elongate, without apicoventral spike, without spe-
cial enlarged apical spine; peduncular apices of
uropods 1-2 not combed; inner ramus of uropod 1
with marginal spines in one row, either no or some
rami continuously spinose to apex, or bearing ac-
cessory nails, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary; article 2 of outer ramus carry-
ing 2 medium to long apical setae. Telson ordinary,
with only 1-2 apical spines, or 2 additional setae on
each lobe plus setules.
DESCRIPTION.?Rostrum fully developed to obso-
lescent. Fuzz on article 1 of antenna 1 in male pres-
ent; calceoli on male primary flagellum of antenna
1 present. Calceoli on article 5 of male antenna 2
192 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
present; flagellum in male with calceoli. Prebuccal
parts ordinary, poorly separated from each other,
upper lip dominant. Right lacinia mobilis bifid or
simple, flabellate or thin; mandibular palp medium
to thin, article 1 short to slightly elongate, article 2
usually without outer setae, apex of article 3
oblique. Lower lip bearing cones. Outer plate of
maxilla 1 with 9?11 spines, one spine especially
thickened. Inner plates of maxilliped thick, ordi-
narily setose. Gills present on coxae 2-7. Coxae 2-4
without special anterodorsal humps. All posterior
spines on article 6 of pereopods 1-2 thick and stiff,
midapical spine or seta usually present. Article 2
of pereopod 5 without facial setae. Peduncle of uro-
pod 1 with dorsolateral spines confined apically,
medial spines widely spread; peduncle of uropod 2
with only one medial spine or setule confined
apically or spines widely spread. Peduncle of uro-
pod 3 lacking extra subapical setae or spines. Tel-
son with ordinary pair of midlateral or dorsal set-
ules on each side.
TYPE-SPECIES.?Birubius panamunus Barnard
and Drummond, 1976 [originally designated].
COMPOSITION.?36 species (in addition to the type
species), all from southern Australia, all described
herein.
RELATIONSHIP.?Birubius differs from Ponthar-
pinia Stebbing in the nontriturative molar bearing
semiarticulate spines and in the small gnathopod
2, among numerous other characters.
Externally, Birubius is difficult to separate from
the generally similar Brolginae but all species have
4 or more spines on the molars whereas the Brolgus
group has only 3. Unlike various members of the
Brolginae, species of Birubius always lack combs
on the peduncles of uropods 1-2, the posterior
spines on article 6 of pereopods 1-2 are never thin
and seta-like, gnathopod 2 is as small as gnathopod
1, article 5 is free, never cryptic, and the other
unusual modifications of gnathopods seen in cer-
tain members of the Brolgus group are absent.
Birubius usually has 4 (rarely 3) setae on the inner
plate of maxilla 1 whereas the Brolginae usually
have only 2. The outer plate of maxilla 1 lacks the
extra midapical cusp or setule of the Brolgus
group. Article 2 of antenna 1 and article 5 of
antenna 2 are always fully developed in Birubius
whereas they usually are reduced in the Brolginae.
The facial spines on article 4 of antenna 2 always
occur in 2 or more fully developed rows whereas
the Brolgus group has these usually in one main
row. The distalmost posterior setule on the dactyl
of pereopods 1-2 in the Brolginae is generally more
distal and more weakly marked than in Birubius.
Article 2 of pereopod 5 in Birubius always bears 2
facial ridges in contrast to the Brolgus group bear-
ing only one. Epimeron 3 of Birubius is rarely as
simple as that of the Brolgus group and articles
4-5 of pereopods 3 and 4 are usually very slightly
to much broader than in the Brolgus group.
From Parharpinia Stebbing, Birubius differs in
the absence of supernumerary dorsal spination or
setation on the telson, lacks the special apicomedial
spine on the peduncle of uropod 1, has a longer
apical nail on the dactyl of the maxilliped, usually
lacks any indication of an ensiform antenna 2, and
never has fully developed posterior setae on epi-
mera 1-2 as seen in certain phases or taxa of Par-
harpinia. Article 2 of pereopod 3 in species of
Birubius usually is untapered though a few species
show faint suggestions of this condition, whereas
this article in Parharpinia tapers strongly and lacks
a posteroventral lobe and furthermore is heavily
setose posteriorly. Article 2 of pereopod 5 in Biru-
bius bears 2 facial ridges whereas Parharpinia bears
one.
Birubius differs from Protophoxus in most of
the same characters as it does from Parharpinia
including those items concerning pereopod 3, tel-
son, uropod 1 (but in Protophoxus the special spine
is lateral), antenna 2, and furthermore lacks a comb
on the peduncle of uropod 2.
Trichophoxus K. H. Barnard and Tipimegus,
new genus, have distinctive molars, mitellid hands
on the gnathopods, lack posteroproximal setae on
article 5 of pereopods 1-2, and have a miniaturized
pereopod 5 with enlarged article 3.
Birubius is most closely similar to the hypothet-
ical immediate ancestor, Cunmurra, new genus,
from which Birubius differs only in the small
gnathopod 2 and enlarged outer plate of the max-
illiped. Cunmurra would not appear to be on a
direct line from a primitive ancestor to Birubius
because of that small plate on the maxilliped.
In this genus the facial spine formula on article
5 of antenna 2 includes clearly apicofacial spines
near or disjunct from the ventrodistal spine group.
NUMBER 245 193
The Evolutionary System in Birubius
FIGURE 82
Birubius is the largest known genus of Phoxo-
cephalidae. Many more new species probably will
be discovered in Australia. Eventually other genera
may be split away from the known 37 species; we
indicate some of these places in the discussions of
relationship among the many species. In the pres-
ent context of our knowledge, however, the com-
position of the genus has been constrained tightly.
The species presented here are arranged in an
evolutionary system with progression away from
what is considered to be the most primitive state,
that represented by B. panamunus in which several
characters are reminiscent of the primitive genus
Pontharpinia. The characters include fully devel-
oped right lacinia mobilis, large tooth and facial
setae on epimeron 3, setose coxa 4, unmodified
head, broad articles of pereopods 3^1, and the
presence, in a reduced state, of continuous dorsal
spination on one ramus of uropods 1-2 as marked
by the presence of accessory apical nails. Other
characters of Pontharpinia, such as strongly devel-
oped ventral setae on article 2 of pereopod 5 are
not found in B. panamunus but do occur in sev-
eral species of Birubius in which other more funda-
mental characters have departed from the pana-
munus model. Such replications of primitiveness
suggest that Birubius may be polyphyletic and that
further divisions will become necessary once more
species have been found to demonstrate the strength
of the various clusters.
The infrageneric evolutionary system in Birubius
is based on: the successive loss of complexity on
epimeron 3, on the right lacinia mobilis, on the
apex of the inner ramus on uropod 1; the loss of
setae on coxa 4 and basofacially on uropod 1; the
decrease in stoutness of pereopods 3?4; the increas-
ing constriction of the rostrum; and the elongation
of the wrists on the gnathopods. Most of the spe-
cies have only 3 teeth on the right mandibular
incisor, few retain long ventral setae on article 2
of pereopod 5, all but one have lost all evidence
of an ensiform process on antenna 2, the second
article of the outer ramus on uropod 3 decreases
in size, the medial and lateral spination on the
peduncles of uropods 1-2 becomes more simplified,
setation is reduced on the mandibular palp, and
spination on article 6 of pereopods 1-2 is reduced
by stages in the evolutionary sequence.
The following characters appear to be increasing
in complexity in the evolutionary scheme: number
of molarial spines, complexity of cuticular orna-
mentation, apical broadening of coxa 1 with con-
comitant decrease in setation, broadening or trape-
zoidal extension of coxa 4, increasing length of
article 2 on antenna 1 with concomitant decrease
in setation, increase in size and facial spination of
article 5 on antenna 2, increasing number of facial
ridges on pereopods 3-5, increasing apical digita-
tion on pereopod 5, and loss of articulation on the
urosome with increase in development of dorsal
processes (very weakly seen).
Key I to Birubius is mainly based on this scheme
of evolutionary flow through the genus. The spe-
cies are arranged generally in the same sequence,
although for each cluster of species we pick a good
model which may or may not be the most primitive
in the cluster and thus may be placed forward out
of sequence. The sequence is shown in Figure 82,
in which the lines and arrows connecting clusters
demonstrate the flow of character modification
rather than indicating precise descent.
Keys to the Species of Birubius (Females)
KEY I (Phyletic)
1. Right lacinia mobilis bifid 2
Right lacinia mobilis simple KEY C
2. Distal branch of right lacinia mobilis complex, either subbifid or toothed KEY A
Distal branch of right lacinia mobilis simple, pointed KEY B
KEY A
1. One or more rami of uropods 1-2, primarily inner ramus of uropod 1, with 1-2 apical acces-
sory nails, hence ramus continuously spinose to apex 2
194 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
No ramus of uropods 1-2 bearing apical accessory nails ' ?
2. Coxa 4 with long ventral setae *
Coxa 4 lacking long ventral setae *
S. Tooth of epimeron 3 large, full face with oblique-horizontal row of setae plus ventral
row ' ? *? ^tunmmuj
Tooth of epimeron S small, full face with 0-3 irregularly set setae not in even row plus
ventral row *
4. Some ventral setae on epimeron 2 in posterior half of face, posteroventral angle of epimeron
2 unprotuberant, urosomite 1 with long lateral row of setae 2. B. torus, new species
FIGURE 82.?Evolutionary pattern in Birubius (numbers indi-
cate species as captioned in the text; 0 ? zero; character sym-
bols are identified at bottom right; arrows indicate general
flow of evolution from primitive level in upper left to most
advanced level in lower right; seven boxes, A-G, surround
species in congruently similar grades of evolution).
Box A bears the most primitive species, with fully flabel-
late, bifid right ladnia mobilis and accessory apical nails on
the rami of uropods 1-2. The head is fully rostrate, and
species 1, the most primitive, bears the enlarged postero-
ventral tooth on epimeron 3. Species 1-3 (left of the dashed
vertical subdivisional line) are species with setose coxa 4,
while species 4-9 (to the right of the dashed line), lack those
setae. Species 2-3 are characterized by loss of facial setae on
epimeron 3, reduction of its tooth, and modification in shape
of coxa 4. Species 4-5 are characterized by disarray of facial
setae on epimeron 3 and reduction in its tooth. Species 6
has epimeron 3 in primitive condition but ventral setae are
lost and basofacial setae on uropod 1 are reduced. Species
7-8 have lost facial setae on epimeron 3 but retain the
ventral setae. Species 9 is characterized by anterior loss of
ventral setae on epimeron 3 and narrowed rostrum.
Box B contains species 10, an offshoot of the Box A group,
in which the right lacinia mobilis has become thin and
nonflabellate but accessory spines on the rami of uropods
1-2 are maintained; species 10 grades into the Box D group
where the accessory spines have also been lost; however the
setation on epimeron 3 has become too simplified for species
10 to be considered an intergrade to the species of Box D.
Box C contains species 11-15, generally similar to those of
Box A but in which uropods 1-2 have lost the accessory
nails on the rami. The model, species 11, has epimeron 3
somewhat similar to species 9 but all species of Box C have
setae on coxa 4 and presumably flow directly from species 1.
Species 13 is distinguished in the loss of the posterodorsal
seta on epimeron 3 and the weakly constricted rostrum.
Species 15 is characterized by a partially replicated and com-
plex right lacinia mobilis. Species 14 bears long ventral setae
on article 2 of pereopod 5. The rostrum becomes somewhat
constricted in the higher numbered species of this group.
Box D contains species 20-27, characterized, like those of
Box B, by the nonflabellate but bifid right lacinia mobilis
but, unlike the species in Boxes A and B, showing the ab-
sence of accessory apical nails on the rami of uropods 1-2.
The model, species 20, has scattered facial setae on epimeron
3, but is considered to be primitive because of the absence
of posteroproximal spines on article 5 of pereopods 1-2. All
other species of Box D bear those spines. Species 22, 23, 24,
26, and 27 have elongate article 5 of gnathopod I. Species
22-23 have facial setae on epimeron 3 organized into a row.
Species 25-27 have reduced basofacial setae on uropod 1.
Species 24 has a very special setation pattern on epimeron 3,
with deeply submarginal posterior setae. In addition, species
20-23 have short anterior facial ridge on female pereopod 5
(symbolized by a star) and species 22-23 have slightly elon-
gate article 2 on outer ramus of uropod 3 (designated by +).
It also should tie noted that on epimeron 3 specie* 20 lucks
ventral setae, specie* 21 and 24 bear several ventral setae,
species 22-23 l>ear only one long ventral seia. species 2.r>-27
bear only tiny posteroventral setule* in notches, and that on
epimeron 2 in females, species 22. 23. 26. 27 have facial setae
crowded forward.
Box E contains species 16-18 with primitive epimeron 3
but with highly advanced mandible bearing an almost sim-
ple right lacinia mobilis lacking anything but a cusp to
represent the distal branch and with unusual, enlarged,
smooth raker spines lacking intercalated plusetules. The
epimeral tooth and strong ventral setation on article 2 of
pereopod 5 are reduced in species 18.
Box F contains species 31 alone. This species combines a
simple right lacinia mobilis with the presence of accessory
nails on the rami of uropods 1-2 plus very thin rostrum,
lack of setae on coxa 4 and reduced basofacial setation on
uropod 1. It forms an excellent bridge between Boxes A
plus C and G.
Box G contains all other species, probably of diverse origin
as shown by the arrows. All carry the simple right lacinia
mobilis and lack accessory apical nails on uropods 1-2, ex-
cept that species 32-37 bear a setule in place of the accessory
apical nail. Species 19 is conceived to flow from Box E where
the distinctions between right lacinia mobilis and epimeron
3 are minor but species 19 has lost setae on coxa 4 and the
setae on coxa 1 are reduced to 2, like species 34-36.
Species 28-29 have their nearest ancestral grade in species
11 as evidenced by epimeron 3, uropod 1, and coxa 4. Species
30, with its unusual epimeron 3 appears to be descendant
from a grade represented by species 24 with similar epi-
meron 3; but species 30 has the simple right lacinia mobilis.
All other species of Box G, 32-37, have lost setae on coxa
4, have the extremely narrowed rostrum and a reduction of
setae on coxae 1-3; on those coxae, species 32-33 have 3
setae, species 34-37 have 2 setae. Species 34-37 retain a
posterodorsal setule on epimeron 3 whereas species 32-33 do
not. Species 34-35 have long ventral setae on article 2 of
pereopod 5. Species 37 is the ultimate evolute in its ex-
tremely reduced rostrum.
NUMBER 245 195
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
No ventral setae of epimeron 2 in posterior half of face, posteroventral angle of epiraeron
2 protuberant, urosomite 1 naked laterally 3- B. nammuldtu, new species
5. Epimeron 3 with more than 2 fully facial setae 6
Epimeron 3 with one or fewer fully facial setae 9
6. Epimeron 3 with ventral setae '
Epimeron 3 naked ventrally 8
7. Mandibular molars with disjunct spine very close to other spines, see descriptions for
minute characters 4 B. myallus, new specie*
Mandibular molars with disjunct spine highly remote from other spines, see description for
minute characters 5. ?. kareus, new species
8. Rostrum weakly constricted, main spine on article 5 of pereopods 1-2 reaching apex of
article 6 in flexed position (male) 6. B. apart, new species
Rostrum unconstricted, main spine on article 5 of pereopods 1-2 falling well short of apex
on article 6 in flexed position (female) 6. B. apart, new species
9. Article 5 of gnathopods 1-2 elongate, epimeron 3 with midposterior setules 10
Article 5 of gnathopods 1-2 short, epimeron 3 lacking midposterior setules
9. B. muldarptts, new specie*
10. Tooth on epimeron 3 of medium size, cuticular setules surrounded by alate plates
7. B. cartoo, new species
Tooth on epimeron 3 small, cuticular setules lacking alatc plates. . 8. B. thalmus, new species
11. Right lacinia mobilis normal, uropod 1 bearing t>av>faci;il setae 12
Right lacinia mobilis with additional accessory proximal bifid process, uropod 1 lacking
basofacal setae 15. B.boolem, new species
Right lacinia mobilis normal, uropod 1 lacking basofacial setae (and article 2 of pereopod
5 with one slightly elongate ventral setulc, inner ramus of uropod 1 with one enlarged
spine, epimeron 3 with posterodorsal setulc, coxa 1 expanded distally, article .r> of pereo-
pods 1-2 with posteroproximal spines, epimeron 3 with only posterior and facial seiae
[these comments provided so as to follow through couplets 12-15 to check this species])
(see KEY B) 26. B. kinkus, new species
12. Article 2 of pereopod 5 lacking elongate ventral setae or with only one such seta 13
Article 2 of pereopod 5 bearing many long ventral setae 14. 0. karobrani, new species
13. Epimeron 3 with posterodorsal setule, spines on inner ramus of uropod 1 of normal size,
coxa 1 expanded distally 14
Epimeron 3 lacking posterodorsal setule, spines on inner ramus of uropod 1 enlarged, coxa
1 unexpanded distally 13. B. chintoo, new species
14. Article 5 of pereopod 2 bearing posteroproximal spine, epimeron 2 not protuberant in
female 12. B. wiraktu, new species
Article 5 of pereopod 2 lacking posteroproximal spine, epimeron 2 protuberant in
female 15
15. Epimeron 3 bearing ventral setae, lacking facial setae, posterior setae short and sparse
11. B. mayamayi, new species
Epimeron 3 lacking ventral setae, bearing facial setae, posterior setae long and numerous
(see KEY B) 20. B. gambodeni, new species
KEY B
1. Inner ramus of uropod 1 with 2 apical accessory nails 10. B. gattangus, new species
Inner ramus of uropod 1 lacking accessory nails 2
2. Distal branch of right lacinia mobilis so small as to form only vestigial cusp on main
branch 3
Distal branch of right lacinia mobilis large, fully formed 5
3. Tooth of epiraeron 3 small 18. B. quearus, new species
Tooth of epimeron 3 large 4
4. Article 2 of pereopod 3 rounded dorsoposteriorly 16. B. babancekus, new species
Article 2 of pereopod 3 with strong dorsoposterior cusp 17. B. gelarus, new species
5. Pereopods 1-2 lacking posteroproximal spine(s) on article 5 (see KEY A)
20. B. gambodeni, new species
Pereopods 1-2 bearing posteroproximal spine(s) on article 5 6
6. Epimeron 3 with 1-3 long ventral setae 7
Epimeron 3 lacking long ventral setae, occasionally with short posteroventral setules 10
NUMBER S45 197
7. Article 5 of gnathopods 1-2 short, facial setae on epimeron S scattered
21. B. maamus, new species
Article 5 of gnathopods 1-2 elongate, facial setae on epimeron 3 either organized into an
even row or absent 8
8. Epimeron S with numerous posterior and ventral setae, no setae fully facial, uropod 1
basofacially setose (5 +), setae on epimeron 2 widely spread 24. B. bateit
Epimeron 3 with only few short posterior setae, only one ventral seta, face with long row
of setae, uropod 1 poorly setose basofacially (0-2), setae on epimeron 2 crowded forward . . 9
9. Basofacial seiae of uropod 1 ?= 0, urosomite 1 articulation line well developed, inner
ramus of uropod 2 with one dorsal spine, and see discussion of these species
22. B. lowarmus, new species
Basofacial setae of uropod I ? 2, urosomite 1 articulation line weak, inner ramus of uropod
2 with 2 dorsal spines, and see discussion of these species . . . . 23. B. kyeenuu, new species
10. Article 1 of mandibular palp slightly elongate, article 5 of pereopods 1-2 with one postero-
proximal spine, (article 5 of gnathopod 1 weakly elongate) .. 27. B. munggai, new species
Article 1 of mandibular palp short, article 5 of pereopods 1-2 with 2 posteroproximal
spines, (article 5 of gnathopod 1 variable) 11
11. Article 5 of gnathopod 1 ordinary, inner ramus of uropod 1 with 2 dorsal spines
25. B. kokorus, new species
Article 5 of gnathopod 1 elongate, inner ramus of uropod I with one dorsal spine (see
KEY A) 26. B. kinkus, new species
KEY C
1. Epimeron 3 with giant posteroventral tooth 19. B. narus, new species
Epimeron 3 with small or no tooth 2
2. Coxa 4 with long ventral setae 3
Coxa 4 lacking long ventral setae 4
3. Head evenly ovate in dorsal view, cuticle relatively simple, see description
28. B. ularitus, new species
Head with narrowed rostrum and sinuous lateral margins dorsally, cuticle complex, see
description 29. B. eleebanus, new species
4. Uropod 1 with 5+ basofacial setae, epimeron 3 with vertical submarginal row of posterior
setae, posterior margin unnotched 30. B. jirrandus, new species
Uropod 1 with 0-1 basofacial seta, epimeron 3 lacking special vertical row of posterior
setae, often with scattered posterior setae or setules in marginal notches 5
5. Epimeron 3 with facial setae 6
Epimeron 3 lacking facial setae 7
6. Rostrum elongate, wrists of gnathopods short, inner ramus of uropod 1 with accessory
apical nails 31. B. yorlunus, new species
Rostrum short, wrists of gnathopods elongate, inner ramus of uropod 1 lacking accessory
nails 32. B. eake, new species
7. Coxae 1-3 with more than 2 elongate ventral setae, epimeron 3 lacking fully posterodorsal
setule 33. B. kabbtdinus, new species
Coxae 1-3 with only 2 elongate ventral setae, epimeron 3 with fully posterodorsal setule.. 8
8. Each lobe of telson with 2 apical spines, rostrum heavily reduced in length
37. B. wulgaru, new species
Each lobe of telson with one apical spine, rostrum very narrow but elongate 9
9. Article 2 of pereopod 5 with few or no long ventral setae, cuticle lacking villi
"36. B. maldus, new species
Article 2 of pereopod 5 heavily setose ventrally, cuticle villose 10
10. Outer ramus of uropods 1-2 with long gap between apex and first apical spine, peduncle
of uropod 1 with 2 apicolateral spines 34. B. taldeus, new species
Outer ramus of uropods 1-2 with spination continuous dorsally to apex, peduncle of uro-
pod 1 with only one apicolateral spine 35. B. yandus, new species
KEY II (Gradal and Identificatory)
1. Inner ramus of uropod 1 with apical accessory nail 2
Inner ramus of uropod 1 lacking accessory nail 13
SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY
198
2. Coxa 4 with long ventral setae
Coxa 4 lacking long ventral setae
3. Tooth of epimeron 3 large, face with long even row of setae I B . panamunus, new species
Tooth of epimeron 3 small, face lacking any but 1-2 scattered setae 4
4. Urosomite 1 with long row of lateral setae 2. B. lorus, new specie*
Urosomite 1 lacking lateral setal row SB. nammuldus, new species
5. Epimeron 3 with 2 or more ventral setae 6
Epimeron 3 with 0-1 ventral seta '
6. Epimeron 3 with setae fully facial '
Epimeron 3 lacking fully facial setae 9
7. Uropod 1 with 0-1 basofacial seta (right lacinia mobilis simple).. 31 B. yorlunus, new species
Uropod 1 with 3+ basofacial setae (right lacinia mobilis bifid) 8
8. Mandibular molars with disjunct spine close to other spines, see descriptions for other
details 4. B. myallus, new species
Mandibular molars with disjunct spine remote from other spines, see descriptions for
other details 5. B. kareus, new species
9. Epimeron 3 with midposterior setules. article 5 of gnathopod 1 elongate 10
Epimeron 3 lacking midposterior setules, article 5 of gnathopod 1 short
9. B. muldarpm, new species
10. Tooth on epimeron 3 large, cuticular setules surrounded by alate plaques
7. B. cartoo, new species
Tooth on epimeron 3 medium-small, cuticular setules lacking alate plate*
B.B. thalmus, new species
11. Epimeron 3 with strong oblique row of setae on face 12
Epimeron 3 with only one facial seta 10. B. gallangus, new species
12. Rostrum weakly constricted, main spine on article 5 of percopods 1-2 reaching apex of
article 6 (male) 6. B. apart, new species
Rostrum unconstricted, main spine on article 5 of pereopods 1-2 falling well short of apex
of article 6 (female) 6. B. opart, new species
13. Coxa 4 with long ventral setae 14
Coxa 4 lacking long ventral setae 23
14. Article 2 of pereopod 5 with 2 or more elongate ventral setules or setae or with numerous
ventral setules 15
Article 2 of pereopod 5 with ventral setules short, sparse, or absent 18
15. Posteroventral tooth on epimeron 3 small, not longer than other posterior teeth (right
lacinia mobilis fully bifid) 14. B. karobram, new species
Posteroventral tooth on epimeron 3 large, much longer than other posterior teeth (if other
teeth present at all), tooth large in any event 16
16. Tooth of epimeron 3 and ventral setae on article 2 of pereopod 5 of short variety
18. B. quearus, new species
Tooth of epimeron 3 and ventral setae on article 2 of pereopod 5 of elongate variety . . . . 17
17. Article 2 of pereopod 3 rounded posterodorsally 16. B. babaneekus, new species
Article 2 of pereopod 3 with sharp posterodorsal cusp 17. B. gelarus, new species
18. Coxa 1 with anterior and posterior marginal tangents perfectly parallel
15. B. booleus, new species
Coxa 1 with anterior and posterior marginal tangents slightly to strongly divergent 19
19. Right lacinia mobilis bifid 20
Right lacinia mobilis simple 22
20. Epimeron 3 lacking posterodorsal setule, spine(s) on inner ramus of uropod 1 enlarged
13. B. chintoo, new species
Epimeron 3 bearing posterodorsal setule, spines on inner ramus of uropod 1 not enlarged
? 21
21. Article 5 of pereopod 2 with posteroproximal spine(s), epimeron 2 not protuberant postero-
ventrally in female 12. B. wirakus, new species
Article 5 of pereopod 2 lacking posteroproximal spine, epimeron 2 protuberant postero-
ventrally in female 11. B. nuiyomoyi, new species
22. Head evenly ovate in dorsal view, cuticle relatively simple 28. B. ularitus, new species
Head with narrowed rostrum and sinuous lateral margins in dorsal view, cuticle complex
(Figure 170) 29. B. eleebanus, new species
23. Epimeron 3 with very large tooth (right lacinia mobilis simple, coxa 1 with only 2 long
NUMBER 245 199
setae) 19. B. narus, new species
Epimeron 3 with small tooth or none 24
24. Epimeron 3 with single submarginal posterior vertical row of setae, other facial setae
absent 25
Epimeron 3 with or without posterior setae but when present, face bearing other setae, or
only facial setae present 26
25. Right lacinia mobilis bifid 24. B. batrit
Right lacinia mobilis simple 30. B. jirrandus, new species
26. Adult coxa 1 (and usually 2-3) with only 2 long setae 27
Adult coxa 1 with 3 or more long setae SO
27. Rostrum vestigial 37. B. wulgaru, new species
Rostrum thin but elongate 28
28. Article 2 of pereopod 5 with few or no long ventral setae, cuticle lacking villi
36. B. maldus, new species
Article 2 of pereopod 5 heavily setose ventrally, cuticle villose 29
29. Outer ramus of uropod 1 with long gap between apex and apicalmost dorsal spine, peduncle
of uropod 1 with 2 apicolateral spines 34. B. taldeus, new species
Outer ramus of uropod 1 with dorsal spination continuous to apex, peduncle of uropod
1 with only one apicolateral spine 35. B. yandus, new species
30. Right lacinia mobilis simple SI
Right lacinia mobilis bifid SS
31. Gap between apices of rami on uropod 1 and apicalmost spines much greater than between
base to most proximal spines, epimeron 3 with posteroventral notch obsolescent or absent
(only males known) 33. B. kabbulinus, new species
Gap between apices of rami on uropod 1 and apicalmost spines not greater than between
base to most proximal spines, epimeron 3 with posteroventral notch well developed,
marked by tooth in both sexes 32
32. Rostrum elongate, wrists of gnathopods short, inner ramus of uropod 1 often with
accessory apical nail (but occasionally not) 31. B. yorlunus, new species
Rostrum short, wrists of gnathopods elongate, inner ramus of uropod 1 lacking accessory
apical nail 32. B. take, new species
33. Article 5 of pereopods 1-2 lacking posteroproximal spines.... 20. B. gambodeni, new species
Article 5 of pereopods 1-2 bearing posteroproximal spine(s) 34
34. Epimeron 3 with 1-3 long ventral setae 35
Epimeron 3 with 0-3 short posteroventral setules only S7
35. Facial setae on epimeron 3 scattered irregularly 21. B. maamus, new species
Facial setae of epimeron 3 arranged in oblique row 56
36. Basofacial setae of uropod 1 ? 0, inner ramus of uropod 2 with one spine, articulation line
of urosomite 1 well developed and see description 22. B. lowannus, new species
Basofacial setae of uropod 1 = 2 , inner ramus of uropod 2 with 2 spines, articulation line
of urosomite 1 weak, and see discussion in text 23. B. kyeetnus, new species
37. Article 1 on mandibular palp slightly elongate, article 5 of pereopods 1-2 with one postero-
proximal spine (article 5 on gnathopod 1 weakly elongate) .. 27. B. munggai, new species
Article 1 of mandibular palp short, article 5 of pereopods 1-2 with 2 posteroproximal spines
(article 5 of gnathopod 1 variable) 38
38. Article 5 on gnathopod 1 ordinary, inner ramus of uropod 1 with 2 dorsal spines
25. B. kokorus, new species
Article 5 on gnathopod 1 elongate, inner ramus of uropod 1 with one dorsal spine
26. B. kmkus, new species
Aids to Identification in Birubius listed with its assigned number to assist finding
the species in the text.
Besides the keys to the species of Birubius, which
emphasize conditions of the right lacinia mobilis Kind of epimeron 3 (see Figure 83 for graphic identification
and inner ramus of uropod 1, the following lists of classes)
r , , . ? i - j ?<- 1; 1. panamunus; 3, nammuldus.
of character alternatives may assist the identifier
 o K ? ? a .u t , ? u u u IT r
1
 2a, b: 7, cartoo; 8, thalmus; 16, babaneekus; 17, gelarus;
in isolating groups of species in this genus. Each
 18 qUearus.
species with the character in positive degree is 3: 19, narus.
200 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY
FIGURE 83.?Kinds of epimeron 3 of Birubius (see text, page 199).
4: 6, apart.
5: 4, myallus; 5, kareus; 21, maamus.
6: 9, muldarpus.
7: 10, gallangus; 32, eake.
8: 31, yorlunus.
9: 22, lowannus; 23, kyeemus.
10: 20, gambodeni; 25, kokorus; 26, kinkus; 27, munggai.
11: 24, bateiT; 30, jirrandus.
12: 2, /orus; 11, mayamayi; 12, wirakus; 14, karobrani; 15,
booleus; 28, ularitus; 29, eleebanus.
13: 34, taldeus; 35, yandus; 36, maldus; 37, wulgaru.
14: 13, chintoo.
15: 33, kabbulinus (male).
Coxa 4 with long ventral setae: 1, panamunus; 2, lorus;
3, nammuldus; 11, mayamayi; 12, wirakus; 13, chintoo;
14, karobrani; 15, booleus; 16, babaneekus; 17, gelarus; 18,
quearus; 28, ularitus; 29, eleebanus.
Article 2 of pereopod 5 with long ventral setae: 14, Aaro-
brani; 18, babaneekus; 17, gelarus; 34, taldeus; 35, yandus;
37, wulgaru.
Article 5 of pereopods 1-2 with posteroproximal spine(s):
1, panamunus; 2, lorus; 4, myallus; 5, kareus; 9, muldarpus;
10, gallangus; 12, wirakus; 15, booleus; 19, narus; 20,
maamus; 22, lowannus; 23, kyeemus; 24, bateif; 25, kokorus;
NUMBER 245 201
26, kinkus; 27. munggai; 30. jirrandus; 31, yorlunus; 32,
eake; 33. kabbulinus; 34. taldeus; 35, yandus; 36, maldus;
37, wulgaru.
Article 5 of antenna 2 with more than one set of midfacial
spines: 1, panamunus; 3, ttammuldus; 4, myallus; 5, kareus;
9, muldarpus; 11, mayamayi; 12, wirakus; 14, karobrani; 15,
booleus; 16, babaneekus; 17, gelarus; 18, quearus; 20,
gambodeni; 24, bateif; 28, ularitus; 29, eleebanus.
Article 5 of antenna 2 with single set of 3 spines: 7, cartoo;
21, maamus; 23, kyeemus; 25, kokorus (occasionally); 27,
munggai; 30, jirrandus (occasionally 4).
Articles 4-5 of pereopod 4 extremely thin: 11, mayamayi;
14, karobrani; 15, booleus; 16, babaneekus; 17, gelarus; 18,
quearus; 28, ularitus.
Pereopod 3 with facial ridge: I, panamunus; 2, /ortu; 3,
nammuldus; 7, cartoo; 8, thalmus; 10, gallangus; 13,
chintoo; 29, eleebanus.
Coxa 1 almost or fully rectangular: 2, lorus; 3, nammuldus;
6, apari; 7, cartoo; 12, wirakus; 13, chintoo; 15, booleus.
Setae widely spread on coxa 1: 1, panamunus; 2, /orus; 3,
nammuldus; 4, myallus; 7, cartoo; 8, thalmus; 11, maya-
mayi; 14, karobrani; 16, babaneekus; 17, gelarus; 18,
quearus.
Coxa 1 bearing only 2 long ventral setae: 19, narus; 34,
taldeus; 35, yandus; 36, maldus; 37, wulgaru.
Setae on article 2 of antenna 1 highly proximal: 2, nurn-
muldus; 4, myallus (weak); 21, maamus; 23, kyeemus; 24,
bateif; 30, jirrandus; 31, yorlunus; 34, taldeus; 35, yandus;
36, maldus.
Telson with more than one posterior spine on each lobe: 8,
thalmus (extra setule); 20, gambodeni (setae); 37, wulgaru:
(16, babaneekus, exceptional specimens).
Urosomite 1 with lateral row of setae: 2, /orui.
Outer ramus of uropod 2 with accessory nail: 31, yorlunus.
Rostrum vestigial: 37, wulgaru.
1. Birubtus panamunus Barnard and Drummond
FIGURES 84-87
Birubius panamunus Barnard and Drummond, 1976:544-545.
DESCRIPTION OF FEMALE.?Head about 21 per-
cent of total body length, greatest width about 73
percent of length; rostrum scarcely constricted,
broad, exceeding middle of article 2 on antenna 1.
Eyes small to medium, largely occluded with pig-
ment. Article 1 on peduncle of antenna 1 almost
1.5 times as long as wide, about 2.5 times as wide
as article 2, ventral margin with about 6 setules,
unproduced dorsal apex with 3 setules; article 2
about 0.85 times as long as article 1, with ventral
cycle of 6 setae; primary flagellum with 7-8 articles,
about half as long as peduncle, bearing short aes-
thetascs; accessory flagellum with 6-7 articles. Spine
formula on article 4 of antenna 2 = $-4-7 or
1-3-4-6, dorsal margin with notch bearing 4 setae,
ventral margin with 7 groups of 1-2 long to me-
dium setae, one ventrodistal long spine; article 5
about 0.85 times as long as article 4, facial spine
formula = 1-2-2-2 or 1-3-2-2, dorsal margin
naked, ventral margin with 7 sets of single setae
plus occasional short seta, 2 ventrodistal long to
medium spines and setae; flagellum about 0.85
times as long as articles 4-5 of peduncle combined,
with 11 articles. Mandibles with weak to medium
palpar hump; right incisor with 3 teeth and notch;
left incisor with 4 humps in 2 branches; right la-
cinia mobilis bifid, distal branch shorter than
proximal, broad, subbifid or denticulate, proximal
branch simple, pointed, with marginal denticles;
left lacinia mobilis with 4 teeth; right rakers 7-9;
left rakers 8-10; molar in form of bulbous hump
demarcated mainly by spines, right and left molars
with 4-5 primarily long spines, plus one short
spine weakly disjunct or not; palp article 1 short,
article 2 with one long inner apical seta and 2?4
other shorter inner setae, and 1-4 outer facial setae,
article 3 about 1.4 times as long as article 2, oblique
apex with 10 spine-setae, basofacial formula = 1-2
or 1-3. Inner plate of maxilla 1 somewhat thin,
bearing one medium apical seta, one similar apico-
medial seta, 2 apicolateral slightly shorter setae;
palp article 2 with 5 apical-medial marginal spines
and 4-5 submarginal setae. Plates of maxilla 2
extending subequally, outer slightly broader than
inner, outer with 4 apicolateral setae, inner with
2?4 medial setae. Inner plate of maxilliped with 2
large, thick apical spines, 3 apicofacial setae, 4-5
medial setae; outer plate with 7-9 medial and api-
cal spines, $-4 apicolateral setae; palp article 1
with apicolateral seta, article 2 with 3 widely
spaced apicolateral setae, medial margin of article
2 moderately to weakly setose, article 3 protuber-
ant, with 3-4 facial setae, 2 lateral setae, nail of
article 4 medium, with 2 accessory setules. Coxa 1
scarcely expanded distally, anterior margin straight;
main ventral setae of coxae 1-4 = 17-15-11-8, pos-
teriormost seta of coxae 1-2 shortened; anterior
and posterior margins of coxa 4 weakly divergent,
posterior margin oblique, convex, posterodorsal
corner sharp, posterodorsal margin long, concave,
width-length ratio of coxa 4 = 14:17. Long poste-
rior setae on article 2 of gnathopods 1-2 and pe-
reopods 1-2 = (4-6)-(7-9)-(4-5)-7, long anteriors =
(4_7)_(10-18)-7-0, short anteriors = l-(2-3)^i-
(1-3), no others. Gnathopods ordinary; width ratios
202 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s
FIGURE 84.?Birubius panamunus, new species, holotype, female "h," 6.5 mm (a ? female "a,1
4.7 mm; b = female "b," 5.5 mm; v ? male "v," 4.6 mm; u; ? female "w," 5.2 mm).
NUMBER 245 203
FIGURE 85.?Birubius panamunus, new species, holotype, female "h," 6.5 mm (v =? male "v," 4.6
mm; w ? female "w," 5.2 mm).
204 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
yP5 yAl
FIGURE 86.?Birubius panamunus, new species, holotype, female "h," 6.5 mm (v = male "v," 4.6
mm; w = female "w," 5.2 mm; y ?= male "y," 6.1 mm).
NUMBER 245 205
W
FIGURE 87.?Birubius panamunus, new species, male "y," 6.1 mm.
206 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
of articles 5-6 on gnathopods 1-2 = 30:31 and
31:31, length ratios = 71:60 and 71:58; palmar
humps small, palms oblique; article 5 of gnathopod
1 elongate, ovate, posterior margin rounded-flat;
article 5 of gnathopod 2 elongate, ovate, posterior
margin rounded-flat. Pereopods 1-2 similar; facial
setae formula on article 4 = 5-8 and 5-8, on article
5 = 6-8 and 6-10; main spine of article 5 extend-
ing to M. 100 + on article 6, article 5 with 2-3
proximoposterior spines, spine formula of article
6 = 4 and 6 or 5 and 6 or 5 and 7 plus middistal
seta; acclivity on inner margin of dactyls of pe-
reopods 1-2 weak, emergent setule long, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule-
seta formula = 9-3-1. Articles 4-5 of pereopods 3-4
broad, facial spine rows moderately developed, fa-
cial ridge formula on article 2 of pereopods 3-5 =
1-2-2; width ratios of articles 2, 4, 5, 6 of pereo-
pod 3 = 44:42:40:23, of pereopod 4 = 66:39:27:16,
of pereopod 5 = 71:24:19:9, length ratios of pereo-
pod 3 = 81:34:37:35, of pereopod 4 = 90:53:45:49,
of pereopod 5 = 98:25:22:21; article 2 of pereopod
5 exceeding apex of article 4; medial apex of arti-
cle 6 lacking digital processes. Posteroventral cor-
ner of epimeron 1 rounded-quadrate, posterior
margin weakly convex, setule above corner, antero-
ventral margin with 6-10 medium setae, postero-
ventral face with 4 medium setae, posterior pair
set vertically; posteroventral corner of epimeron
2 weakly protuberant, guarded by setule sinus,
posterior margin convex, facial setae = 12-14, pos-
teriormost pair set vertically; posteroventral corner
of epimeron 3 rounded, with large tooth, posterior
margin weakly concave, one small setule in crotch,
ventral margin with 8-10 setae evenly spread, face
with horizontal row of 7-9 setae in middle, epi-
meron 3 with weak rugosities at posterodorsal
imagination. Urosomite 1 naked, articulation line
incomplete; urosomite 3 weakly protuberant dor-
sally. Rami of uropods 1-2 with articulate en-
larged apical nails, inner ramus of uropod 1 with
two accessory nails, outer ramus of uropod 1 with
5-7 (rarely 3-8) dorsal spines, inner with 2 (rarely
3), outer ramus of uropod 2 with 3-4 dorsal spines,
inner with one dorsomedial spine; peduncle of
uropod 1 with 3-4 apicolateral spines, and 3-4
basofacial setae, medially with 4-5 marginal setae
and spines, apicalmost enlarged; peduncle of uro-
pod 2 with 6-7 dorsal spines, medially with one
apical spine. Peduncle of uropod 3 with 8-9 ven-
tral spines, dorsally with one lateral spine, 1-2
medial spines; rami submasculine, inner extend-
ing to M. 100+ on article 1 of outer ramus, apex
with 2 setae, medial and lateral margins with
1 and 4 setae, article 2 of outer ramus short, 0.18,
bearing 2 long setae, apicomedial margin of article
1 with 2 setae in tandem, lateral margin with 4
acclivities, spine formula = 1-1-1-1-1, setal for-
mula = 1-1-1-1-1. Telson short, length-width
ratio = 13:15, almost fully cleft, each apex wide,
rounded, lateral acclivity deep, narrow, (tearing
ordinary lateral setule, spine next medial of length
equal to setule, midlateral sctules diverse. Cuticle
with ordinary bulbar setules, emergent setules short.
DESCRIPTION OF MALE.?Rostrum slightly nar-
rower than in female. Article 2 of antenna 1 with
6 ventral setae; primary flagellum with 7-9 articles,
one calceolus each on articles 1-5, aesthetascs
strongly developed. Facial spine formula on article
4 of antenna 2 = 3-4-4 or 3-4-5 or 3-5-5 or
3-1-4-4, on article 5 = 2-2 or 2-1, article 5 with
4-5 dorsal sets of male setae and 4 calceoli, ventro-
distal apex with one thin spine and 2 setules, fla-
gellar formula = (28-36), 2-3^1-5, 7, 9 . . . penulti-
mate. Basofacial setal formula on article 3 of mandi-
bular palp = 2-2 or 3-2. Coxa 4 more elongate than
in female; ventral setal formula of coxae 1-4 = 15-
(12-14)-8^i. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 3-5-3-5, long
anteriors = 0-4-7-(4-5). Facial and setal-spine for-
mulas of pereopods 1-2 on article 4 = 6-7 and 6,
on article 5 = 4-6 and 7-8, on article 6 = 4 + 5
and 5 + 5 plus middistal seta. Article 2 of only
pereopods 4-5 narrower than in female, article 6
of pereopods 3 and 5 and articles 5-6 of pereopod 4
thinner and more elongate than in female; spines
and setae shortened on articles 2-3 of gnathopods
1-2 and pereopods 1-2 and 5. Coxa 6 with poste-
rior lobe deepened. Article 5 of pereopod 5 with
special posterodistal spine. Epimera 1-3 broadened;
setal formulas, epimeron 1 anteroventral = 10-14,
epimeron 2 facial = 6-11, (anterior = 1-3, absent
in female), epimeron 3 facial = 4-7, ventral = 3-7.
Urosome elongate, narrowed, declivity of fused
urosomites 1-2 sharpened, articulation strongly
marked. Spine formulas of uropods, uropod 2 pe-
duncle dorsal = 10-11, dorsal spines on outer ramus
of uropod 1 = 5 , ventral spines on peduncle of
uropod 3 = 7, spine formula on article 1 of outer
ramus = 1?0?1?1?1?1?I, setal formula = 1-2-1-1-
NUMBER 243 207
1-1-1 or 1-1-1-1-1-1-1. Telson elongate, distal
spines shortened.
OBSERVATIONS (male).?Male "y," 6.1 mm:
Though unusually large, is immature, lacking
penes; eyes relatively small; dorsal pleonal ridge
poorly defined; rami of uropod 3 underdeveloped;
peduncle of uropod 1 with only 3 basofacial setae,
of uropod 2 with only 7 dorsal spines; spine for-
mula on article 4 of antenna 2 = 1-3-3-5, on arti-
cle 5 = 2-2; basofacial formula of article 3 on man-
dibular palp = 1-1; inner plate of maxilla 1 with
only 2 stout setae, one palp bearing extra apical
spine; article 5 of pereopods 1-2 with 3 proximo-
posterior spines; epimera poorly setose.
Male "v" 4.6 mm: Facial spines on article 4 of
left antenna 2 abnormal = 3-1-4-4, erratically ar-
ranged; article 5 devoid of facial spines; right an-
tenna 2 normal.
ILLUSTRATIONS.?The inner plate of the maxilli-
ped bears one small, simple, apicolateral and dor-
sal seta not shown; dorsomedial spine on inner
ramus of uropod 2 hidden, not shown; anterior
marginal setae on epimeron 3 of male omitted;
right and left sides of illustrated male head dis-
tinctive owing to bilateral deformation; article 4
of antenna 2 on male "v" not flattened but article
5 fully flattened.
HOLOTYPE.?NMV, female "h," 6.5 mm.
TYPE-LOCALITY.?PPBES 337, 11 Mar 1971, Port
Phillip Bay, Victoria, Australia, 8 m, fine sand.
VOUCHER MATERIAL (all illustrated).?Type-
locality, female "w," 5.2 mm; PPBES 919/5, male
"v," 4.6 mm; PPBES 919/4, female "b," 5.5 mm;
PPBES 901/4, female "a," 4.7 mm; CPBS 31E/1,
male "y," 6.1 mm.
RELATIONSHIP.?Birubius panamunus (1) is con-
sidered to be the primitive model of Birubius in
which numerous characters bear weak resemblance
to more primitive phoxocephalids such as Pon-
tharpinia. These characters are the fully broad
rostrum, the fully developed right lacinia mobilis,
the large tooth on epimeron 3, at least a few facial
setae on the epimera, basofacial setae on the pe-
duncle of uropod 1, long setae on coxa 4, the pres-
ence of accessory nails, at least on the inner ramus
of uropod 1, thus reflecting remnants of continuous
dorsal spination seen in all rami of uropods 1-2
in Pontharpinia, and reasonably broad articles 4-5
of pereopods 3-4.
Another possible model is B. mayamayi
 v'll) but
in that species the epimeral tooth and setae are
reduced. Birubius gambodeni (20) lacks setae on
coxa 4, lacks the accessory nails on uropod 1 and
has a smaller tooth on epimeron 3.
MATERIAL.?PPBES, 30 samples from 10 stations
(139); CPBS, 2 samples from 2 stations (3); SBS,
2 samples from 2 stations (2).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 6-22 m, sand, silty sand, clay. New
South Wales: off Malabar, 66 m.
2. Birubius lorus, new species
FIGURES 8&-90
DESCRIPTION OF FEMALE.?Head about 17 per-
cent of total body length, greatest width about 65
percent of length; rostrum constricted, narrow, al-
most reaching middle of article 2 on antenna 1.
Eyes large, mostly occluded with pigment. Article
1 on peduncle of antenna 1 about 1.3 times as long
as wide, about 2.2 times as wide as article 2, ven-
tral margin with about 9 setules, unproduced dor-
sal apex with 3 setules; article 2 about 0.9 times
as long as article 1, with ventral cycle of 4-5 setae;
primary flagellum with 11 articles, about 1.1 times
as long as peduncle, bearing long aesthetascs; ac-
cessory flagellum with 7 articles. Spine formula on
article 4 of antenna 2 = 1-3-4-4 or 1-3-4-3, dorsal
margin with notch bearing 3 setae, ventral margin
with 5 groups of 1-2 long to medium setae, one
ventrodistal medium spine; article 5 almost 0.85
times as long as article 4, facial spine formula =
1-2, dorsal margin bearing 2 sets of short setae,
ventral margin with 4 sets of 1-2 long to short
setae, 2 ventrodistal long to medium spines and
setae; flagellum about 1.4 times as long as articles
4-5 of peduncle combined, with 12 articles. Man-
dibles with medium palpar hump; right incisor
with 4 teeth; left incisor with 6 teeth; right lacinia
mobilis bifid; distal branch shorter than proximal,
flabellate, subtrifid, denticulate, with one facial
hump, proximal branch simple, pointed, with mar-
ginal denticles or facial humps; left lacinia mobilis
with 4 teeth, middle teeth shortened; right rakers
5-6; left rakers 5-6; molar in form of bulbous
hump, right molar with 5 primarily long spines
plus one spine strongly disjunct, left molar with
10 primarily long spines, no spine disjunct; palp
article 1 short, article 2 with one short inner apical
208 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
/ Mt
FIGURE 88.?Birubius lorus, new species, holotype, female "a," 4.60 mm (u ? male "u," length
unknown).
NUMBER 245 209
seta and 2 other shorter inner setae, article 3 about
1.2 times as long as article 2, oblique apex with
9-10 spine-setae, basofacial formula = 1-1 or 0-1.
Inner plate of maxilla 1 large, bearing cne long
apical pluseta, one similar apicomedial seta, 2 api-
colateral much shorter setae; palp article 2 with 5
apicalmedial marginal spines and 5 submarginal
setae. Plates of maxilla 2 extending subequally,
outer scarcely broader than inner, outer with 2
apicolateral setae, inner with no medial setae. Inner
plate of maxilliped with 3 large, thick apical
spines, 3 apicofacial setae, 4 medial setae; outer
plate with 7 medial and apical spines, one apico-
lateral seta; palp article 1 with apicolateral spine,
article 2 with 4 lateral setae, medial margin of
article 2 moderately setose (naked apically), article
3 scarcely protuberant, with 5 facial setae, 3 lateral
setae, nail of article 4 long, with 2 accessory setules.
Coxa 1 not expanded distally, anterior margin al-
most straight; main ventral setae of coxae 1-4 =
(10?13)? 10?(4?5)?3, posteriormost seta of coxae 1-3
shortened; anterior and posterior margins of coxa
4 strongly divergent, posterior margin very oblique,
convex, posterodorsal corner subsharp, posterodor-
sal margin short, undulant, width-length ratio of
coxa 4 = 9:10. Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 = 3-4-4-
(5-6), long anteriors = 7-(ll-12)-O-0, short ante-
riors = 1-1-4-1. Gnathopods thin; width ratios of
articles 5-6 on gnathopods 1-2 = 28:28 and 29:30,
length ratios = 78:61 and 75:58; palmar humps
small, palms weakly oblique; article 5 of gnatho-
pods 1-2 elongate, ovate, posterior margin rounded-
flat, long. Pereopod 2 slightly stouter than pereo-
pod 1, especially article 4; facial setae formula on
article 4 = 5 and 6, on article 5 = 4 and 6; main
spine of article 5 extending to M. 100 on article 6,
article 5 with one proximoposterior spine; spine
formula of article 6 = 3 and 5 or 4 and 5 plus mid-
distal seta, some spines especially long; acclivity on
inner margin of dactyls of pereopods 1-2 sharp,
produced as tooth, emergent setule long, midfacial
pluseta long. Coxae 5-7 posteroventral setule for-
mula = 1-1-1. Articles 4-5 of pereopods 3?4 of
ordinary width, facial spine rows dense to moder-
ately developed, facial ridge formula on article 2
of pereopods 3-5 = 1-2-2; width ratios of articles
2, 4, 5, 6 of pereopod 3 = 44:42:40:21, of pereo-
pod 4 = 60:38:25:12, of pereopod 5 = 61:23:20:9,
length ratios of pereopod 3 = 77:33:42:40, of
pereopod 4 = 82:49:38:46, of pereopod 5 = 87:
21:20:23, article 2 of pereopod 5 almost reach-
ing apex of article 4; medial apex of article 6
deeply combed and bearing 4 digital processes. Pos-
teroventral corner of epimeron 1 rounded-quad-
rate, posterior margin convex, corner with setule,
anteroventral margin with 3-7 medium setae, pos-
teroventral face with 2 long setae; posteroventral
corner of epimeron 2 rounded-quadrate, with se-
tule, posterior margin convex, facial setae = 9-10,
posteriormost pair set vertically; posteroventral
corner of epimeron 3 weakly and broadly protu-
berant, with setule sinus, posterior margin straight,
with 7 setule notches, ventral margin with 3?4
submarginal setae mainly posterior. Urosomite 1
with lateral horizontal row of 5-11 setae, articula-
tion line complete; urosomite 3 unprotuberant dor-
sally. Rami of uropods 1-2 with articulate enlarged
apical nails, inner ramus of uropod 1 with two ac-
cessory nails, outer ramus of uropod 1 with 3-4
dorsal spines, inner with one, outer ramus of uro-
pod 2 with 1-2 dorsal spines, inner with one dorso-
medial spine; peduncle of uropod 1 with 2 apico-
lateral spines, and 1-3 basofacial setae, medially
with 3 marginal setae and spines, apicalmost an
enlarged spine; peduncle of uropod 2 with 4 dorsal
spines, medially with one large, apical spine. Pe-
duncle of uropod 3 with 5 ventral spines, dorsally
with one lateral spine, one medial spine; rami
masculine, inner extending to M. 100 on article 1
of outer ramus, apex with 3 setae, medial and
lateral margins setose, article 2 of outer ramus
ordinary, 0.17, bearing 2 medium setae, medial
margin of article 1 with 4 setae, lateral margin
with 4 acclivities, spine formula = 0-1-1-1-1, setal
formula = 1-1-1-1-1. Telson long, length-width
ratio = 6:5, almost fully cleft, each apex wide,
rounded, lateral acclivity shallow, weak, bearing
short ordinary lateral setule, spine next medial of
length equal to setule, midlateral setules diverse.
Cuticle with ordinary bulbar setules of varying
sizes closely packed with pipes and horseshoes, sur-
face bearing fine striations in form of linear finger-
print pattern, emergent setules ordinary.
DESCRIPTION OF MALE.?Only known specimen
broken, missing urosome and epimeron 3. Rostrum
narrower than in female. Eyes very large, forming
weak lateral bulges. Article 2 of antenna 1 with 5
ventral setae; primary flagellum with 11 articles,
one calceolus each on articles 1-8, one aesthetasc
210 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
G2t
FIGURE 89.?Birubius lorus, new species, holotype, female "a," 4.60 mm (u - male "u," length
unknown).
NUMBER 245 211
RS
FIGURE 90.?Birubius lorus, new species, holotype, female "a," 4.60 mm (u ?= male "u," length
unknown).
strongly developed on articles 9 and 10; accessory
flagellum with 7 articles. Facial spine formula on
article 4 of antenna 2 = 3-4-1, on article 5 = 2
plus 1-3 accessory setules (varying right and left),
article 5 with 6 dorsal sets of male setae and 2-6
calceoli, ventrodistal apex with 2 thin spines and
one setule; flagellar formula = 23 (broken), 1-2-3?4,
6, 8 . . . 22. Right rakers 6 plus 1-2 rudimentaries;
left rakers 7 plus one rudimentary; right molar
with 7 spines plus one weakly disjunct spine; left
molar with 6 spines and one rudimentary spine,
none disjunct; basofacial setal formula of article 3
212 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
on mandibular palp = 1-2. Inner plate of maxilli-
ped with only 2 apical spines. Coxa 4 like that of
female but smaller in relation to coxa 1; ventral
setal formula of coxae 1-4 = (10-11 )-(7-8)-5-3.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = 1?(1?3)?3?3, long anteriors =
(5-6)-8-0-0, short anteriors = 1-1-3-1, no others.
Articles 4-5 of pereopods 1-2 thinner than in fe-
male, especially pereopod 1. Article 2 of only pereo-
pod 3 narrower than in female, articles 4-5 of pe-
reopod 3 also narrower, article 4 of pereopod 4
slightly broadened; pereopod 5 larger relative to
pereopod 3 than in female. Epimera 1-2 (3 un-
known) broadened; setal formulas, epimeron 1
anteroventral = 5, posteroventral = 2, epimeron 2
facial = 8. Cuticle with mainly bulbar setules and
their rudiments, pipes and horseshoes obsolescent,
striations strong on pereopods 3-5 and epimera.
HOLOTYPE.?Figures herein presented of female
"a," 4.60 mm. (The holotype is designated as the
figures because during placement of parts of that
specimen in its depository vial a few parts of the
holotype of Birubius eleebanns were mixed into
the vial; during attempts to segregate the mixture
the telson and one maxilla 2 of B. lorus were lost;
one maxilla 2 of B. eleebanus may be deposited in
this vial as this mouthpart in the two species is
indistinguishable; the holotype specimen may there-
fore be confounded but designation of the figures
ameliorates the problem. No other specimen is as
desirable as this for designation as a holotype.)
TYPE-LOCALITY.?EBS 363, 10 Jul 1973, Lake
Macquarie, Salt's Bay, New South Wales, Australia,
0.3 m, coarse sand.
VOUCHER MATERIAL.?Type-locality: female "b,"
4.00 mm; female "c" 3.90 mm; female "d," 3.93
mm; male "u," broken, true length unknown
(illus.).
RELATIONSHIP.?The B. lorus-nammuldus (2-3)
group of species is very close to B. panamunus (1)
but differs in the absence of a fully facial oblique
row of setae, in the smaller tooth on epimeron 3,
and in the more trapezoidal coxa 4.
Birubius gallangus (10) has a simple distal
branch on the right lacinia mobilis, ordinary gnath-
opodal hands and lacks long setae on coxa 4.
MATERIAL.?EBS, 4 samples (14).
DISTRIBUTION.?New South Wales, Lake Mac-
quarie, Salt's Bay, 0.3 m, coarse sand.
3. Birubius nammuldus, new species
FIGURES 91 , 92
DESCRIPTION OF MALE.?Head about 17 percent
of total body length, greatest width about 75 per-
cent of length; rostrum slightly constricted, reach-
ing (unbent) middle of article 2 on antenna 1. Eyes
large, mostly clear of pigment but stained ochra-
cerus-rose, ommatidia especially large. Article 1 on
peduncle of antenna 1 about 1.1 times as long as
wide, about 2.5 times as wide as article 2, ventral
margin with about 17 setules, weakly produced
dorsal apex with 3 setules; article 2 about 0.9 times
as long as article I, with ventral cycle of 6 setae;
primary flagellum with 8-9 articles, about 0.55
times as long as peduncle, tearing long aesthetascs
and one calceolus each on articles 1-4; accessory
flagellum with 6 articles. Spine formula on article
4 of antenna 2= 3-4-4, dorsomedial margins of
articles 3-4 fuzzy, ventral margin of article 4 with
6 groups of 1-3 short to medium setae, one ventro-
distal medium spine, article 5 about 1.25 times as
long as article 4, facial spine formula = 2-2, dorsal
margin bearing 7 calceoli and 7 sets of male setae,
ventral margin with 5 widespread setae, 5 ventro-
distal setules, flagellum elongate, flagellar for-
mula - (28-30), 1-5, 7, 9 . . . (27 or 29). Mandibles
with weak palpar hump; right incisor with 3 teeth
and notch; left incisor with 5 humps in 2 branches;
right lacinia mobilis bifid, distal branch much
shorter than proximal, broad, subbifid, denticulate,
proximal branch simple, pointed, with marginal
denticles; left lacinia mobilis with 4 teeth plus one
accessory tooth, middle teeth shortened; right rak-
ers 7 plus one rudimentary; left rakers 7; molar
in form of short protrusion, demarcated mainly by
spines, right and left molars with 3 primarily long
spines, plus one short spine weakly disjunct; palp
article 1 short, article 2 with 2 long inner apical
setae and 3 other shorter to much shorter inner
setae, article 3 as long as article 2, oblique apex
with 9 spine-setae, basofacial formula = 2-2. Inner
plate of maxilla 1 thin, bearing one long apical
pluseta, one shorter apicomedial pluseta, 2 apico-
lateral similar setae; palp article 2 with 4 apical-
medial marginal spines and 4 submarginal setae.
Plates of maxilla 2 extending subequally, outer
broader than inner, outer with 3 apicolateral setae,
inner with 2 medial setae. Inner plate of maxilli-
NUMBER 145 213
s
FIGURE 91.?Birubius nammuldus, new species, holotype, male "a," 4.98 mm.
214
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 92.?Birubius nammuldus, new species, holotype male "a," 4.98 mm (y ? male "y,"
5.07 mm).
NUMBER 245 215
ped with 2 large, thick apical spines, 2 apicofacial
setae, 4 medial setae; outer plate with 7 medial
and apical spines, 3 apicolateral setae; palp article
2 with 3 groups of one each apicolateral setae, me-
dial margin of article 2 weakly setose, article 3
weakly protuberant, with 3 facial setae, 3 lateral
setae, nail of article 4 long, with 2 accessory setules.
Coxa 1 scarcely expanded distally, anterior margin
weakly convex; main ventral setae of coxae 1-4 =
(12-14)-(8-10)-(7-8)-(4-5), posteriormost seta of
coxae 1-2 shortened; anterior and posterior mar-
gins of coxa 4 strongly divergent, posterior margin
oblique, convex, posterodorsal corner rounded, pos-
terodorsal margin ordinary, width-length ratio of
coxa 4 = 3:4; long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = (2-3)-4-3-4,
long anteriors = 6-7-(7-8)-(3-4), short anteriors =
(l-2)-2-(2-3)-(0-l). Gnathopods somewhat thin;
width ratios of articles 5-6 on gnathopods 1-2 =
28:29 and 27:28, length ratios = 69.61 and
67:58; palmar humps ordinary, palms weakly
oblique; article 5 of gnathopods 1-2 elongate,
ovate, posterior margin rounded. Pereopods 1-2
similar; facial setae formula on article 4 = 6 and
5, on article 5 = 6 and 7; main spine of article 5
extending to M. 100+ on article 6, article 5 lack-
ing proximoposterior spine; spine formula of arti-
cle 6 = 5 and 5 (or 6 and 5 on pereopod 2) plus
middistal seta, some spines especially long; accliv-
ity on inner margin of dactyls of pereopods 1-2
obsolescent, emergent setule short, almost fully im-
mersed, midfacial pluseta very short, highly distad.
Coxa 5-7 posteroventral setule formula = 1-2-1.
Articles 4-5 of pereopods 3-4 of ordinary width,
facial spine rows sparse to moderately developed,
facial ridge formula on article 2 of pereopods
3-5 = 1-2-2; width ratios of articles 2, 4, 5, 6 of
pereopod 3 = 44:41:36:22, of pereopod 4 = 60:
35:22:12, of pereopod 5 = 66:22:19:8, length ra-
tios of pereopod 3 = 78:36:34:37, of pereopod
4 = 84:52:46:51, of pereopod 5 = 118:25:23:19;
article 2 of pereopod 5 reaching middle of article 5,
medial apex of article 6 finely combed and bearing
5 digital processes. Posteroventral corner of epi-
meron 1 rounded, posterior margin deeply convex,
with setule, anteroventral margin with 8 short to
medium setae, posteroventral face with 2-3 long
setae, one pair set vertically; posteroventral corner
of epimeron 2 rounded, weakly protuberant,
guarded by dorsal setule sinus, posterior margin
convex, facial setae = 7-12, posteriormost or occa-
sional other pair set vertically; posteroventral cor-
ner of epimeron 3 with setule sinus and medium
tooth, posterior margin convex, with one long seta,
ventral margin with 7 setae evenly spread, face
with 1-2 other setae, epimeron 3 with setule on
posterodorsal margin set in weak notch. Urosomite
1 naked, articulation line complete; urosomite 3
unprotuberant dorsally. Rami of uropods 1-2 with
articulate enlarged apical nails, inner ramus of
uropod 1 with two accessory nails, outer ramus of
uropod 1 with 6 dorsal spines, extending full mar-
gin, inner with 2, outer ramus of uropod 2 with
4 dorsal spines, inner with one dorsomedial spine;
peduncle of uropod 1 with 2-3 apicolateral spines,
and 3 basofacial setae, medially with 4 marginal
setae and spines; peduncle of uropod 2 with 7-9
dorsal spines, medially with one small apical spine.
Peduncle of uropod 3 with 6 ventral spines, dor-
sally with one lateral spinule, one medial spine
and setule; rami masculine, inner extending to M.
100 on article 1 of outer ramus, apex with 2 setae,
medial and lateral margins setose, article 2 of outer
ramus short, 0.14, bearing 2 long setae, medial mar-
gin of article 1 setose, lateral margin with 6 ac-
clivities; spine formula = 0-0-1-2-2-2-2, setal for-
mula = Is?2?1?1?1?1?1. Telson elongate, length-
width ratio = 33:29, not fully cleft, wide, rounded,
lateral acclivity weak, bearing short lateral setule,
spine next medial of length subequal to setule, mid-
lateral setules diverse. Cuticle with sparse ordinary
bulbar setules of varying sizes mixed with pipes,
surface bearing fine striations in form of linear
fingerprint pattern, emergent setules short.
ILLUSTRATIONS.?Inner plate of maxilla 1 not
fully flattened in illustration; eyes of all known
specimens composed of clear ommatidia but dor-
sally jumbled, illustration omitting dorsal omma-
tidia except for outline; holotype with right epi-
meron 3 bearing only setule at midfacial position
of spine.
HOLOTYPE.?WAM, male "a," 4.98 mm.
TYPE-LOCALITY.?WAM 414-73, 2 Sep 1954, Bar-
row Island, Western Australia, presumably neritic.
VOUCHER MATERIAL.?Type-locality: male "y,"
5.07 mm (illus.); male "c," 5.00 mm; male "d,"
4.80 mm. WAM 9561: male "m," 4.97 mm; male
"k," 5.58 mm; male "v," 4.61 mm; male "z," 4.55
mm. Female unknown.
RELATIONSHIP.?This species differs from B. lorus
216 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(2) in the more crowded setae of epimeron 2, the
protuberant posteroventral angle of epimeron 2,
the longer spines on the rami of uropods 1-2, the
absence of lateral setae on urosomite 1, the pres-
ence of a ridge on pereopod 3, a weaker tooth on
the dactyls of pereopods 1-2, and in a sharper and
larger tooth on epimeron 3.
MATERIAL.?WAM, 2 samples (8).
DISTRIBUTION.?Western Australia, from Barrow
Island south of Garden Island, neritic.
4. Birubius myallus, new species
FIGURES 95-97
DESCRIPTION OF FEMALE.?Head about 19 per-
cent of total body length, greatest width about 73
percent of length; rostrum weakly constricted,
broad, reaching middle of article 2 on antenna 1.
Eyes large, clear of pigment. Article 1 on peduncle
of antenna 1 almost 1.5 times as long as wide,
about 2.3 times as wide as article 2, ventral margin
with about 13 setules, weakly produced dorsal apex
with 2 setules; article 2 about 0.8 times as long as
article 1, with ventral cycle of 9 setae; primary fla-
gellum with 11-14 articles, about 0.63 times as long
as peduncle, bearing numerous aesthetascs; accessory
flagellum almost as long as primary, with 9-11 arti-
cles. Spine formula on article 4 of antenna 2 =
l-3-4-(5-7), dorsal margin with notch bearing 3
setae and setule, ventral margin with 10 groups of
1-2 long to short setae, one ventrodistal long spine;
article 5 about 0.8 times as long as article 4, facial
spine formula = 2-2-2-2, 1-2-2-2, 1-2-2-1, or
1-2-2-2-1, dorsal margin naked, ventral margin
with 8 sets of 1-2 long to short setae, 3 ventrodistal
medium spines; flagellum about 0.9-1.1 times as
long as articles 4-5 of peduncle combined, with
11-17 articles. Mandibles with weak palpar hump;
right incisor with 3 teeth and midcusp; left incisor
with 5 humps in 2 branches; right lacinia mobilis
bifid, distal branch shorter than proximal, distal
branch flabellate, subbifid, broad, denticulate,
proximal branch simple, with marginal denticles;
left lacinia mobilis with 4 teeth; right rakers 9-14;
left rakers 11?14(?17); molar in form of elongate
bulbous plaque, right molar with 5-9 primarily
long spines, one short spine weakly disjunct, each
molar with setule; palp article 1 short, article 2
with one medium inner apical seta and 3-4 other
shorter inner setae, and groups of 1 and 3 outer
setae, article 3 about 1.2 times as long as article 2,
oblique apex with 10-11 spine-setae, basofacial
formula = 3-1 or 3-2, occasionally 1-2 outer setae.
Inner plate of maxilla 1 ordinary, especially
pointed, bearing one long apical pluseta, one simi-
lar apicomedial seta, 2 apicolateral much shorter
setae; palp article 2 with 5-6 apicalmedial marginal
spines and, 5 submarginal setae. Inner plate of
maxilla 2 scarcely shorter than outer, of equal
breadth, outer with 4-6 apicolateral setae, inner
with 2-3 medial setae. Inner plate of maxilliped with
2-3 large, thick apical spines, 2-3 apicofacial setae,
5 medial setae; outer plate with 9-10 medial and
apical spines, 5 apicolateral setae; palp article 1
with 1-2 apicolateral setae, article 2 with as many
as 6 groups of 1-3 apicolateral setae, medial margin
of article 2 strongly setose, article 3 slightly protu-
berant, with 6 facial setae, 5 lateral setae, nail of
article 4 long, with 2 accessory setules. Coxa 1
weakly expanded distally, anterior margin weakly
convex; main ventral setae of coxae 1-4 = (10?14)?
(10?14)?(10?14)?0, posteriormost seta of coxa 1
slightly shortened; anterior and posterior margins
of coxa 4 weakly divergent, posterior margin al-
most straight, posterodorsal corner subsharp, pos-
terodrosal margin short, undulant, width-length
ratio of coxa 4 = 13:16. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
(3-4)-(5-8)-6-(7-8), long anteriors = 8-(9-19)-
(5-6)-(3-10), short anteriors = (S-6)-($-4)-($-5)-
(3-10), no others. Gnathopods generally ordinary
but hands somewhat thin; width ratios of articles
5-6 on gnathopods 1-2 = 27:30 and 28:28, length
ratios = 74:55 and 70:56; palmar humps small,
palms oblique; article 5 of gnathopod 1 elongate,
ovate, posterior margin flat, long; article 5 of gna-
thopod 2 elongate, ovate, posterior margin rounded.
Pereopods 1-2 similar; facial setae formula on arti-
cle 4 = 7 and (5-7), on article 5 = 6-7; main spine
of article 5 extending to M. 85 on article 6, article
5 with 2-3 proximoposterior spines; spine formula
of article 6 = 5-4 or 6-5 plus middistal seta; accliv-
ity on inner margin of dactyls of pereopods 1?2
weak, emergent setule long, midfacial pluseta ordi-
nary. Coxae 5-7 posteroventral setule-seta form-
ula = (10_12)-(4-7)-(3-6). Articles 4-5 of pereo-
pods 3-4 ordinary, facial spine rows moderately
developed, facial ridge formula on article 2 of
pereopods 3-5 = 0-2-2, width ratios of articles 2, 4,
NUMBER 245 217
FIGURE 93.?Birubius myallus, new species, holotype, female "a," 9.8 mm (w ?= female "w," 5.4
mm; y ? male "y," 7.6 mm).
218 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 94.?Birubius myallus, new species, holotype, female "a," 9.8 mm (y ? male "y," 7.6 mm).
5, 6 of pereopod 3 = 41:40:34:17, of pereopod 4 =
63:36:27:12, of pereopod 5 = 72=22:20:7, length
ratios of pereopod 3 = 78:33:36:38, of pereopod
4 = 85:53:45:?4, of pereopod 5 = 95:23:22:21;
article 2 of pereopod 5 reaching apex of article
4 or beyond in subadult females, with longer seta
on posteroventral angle (absent in illustration); me-
dial apex of article 6 poorly combed, ragged, bear-
ing 5-6 digital processes. Posteroventral corner of
epimeron 1 rounded or sharply protuberant,
posterior margin almost straight, bearing
setule, anteroventral margin with 8-11 long setae,
posteroventral face with 2-5 medium to long setae,
posterior pair or triad set vertically in subterminal
female; posteroventral corner of epimeron 2 with
small protuberance guarded by sinus (setule present
or absent), posterior margin weakly convex, facial
setae = 8-16, posteriormost pair set vertically or
not; posteroventral corner of epimeron 3 with small
to medium tooth, posterior margin straight or un-
dulant, with setule notches and 1-4 setae, face of
ventral margin with 5^ 6 setae.pointing posteriorly,
evenly spread, face with 2-8 scattered setae; epi-
meron 3 with small seta on posterodorsal margin
set in weak notch. Urosomite 1 with midventral
setule, articulation line almost complete; urosomite
3 weakly protuberant dorsally. Rami of uropods
1-2 with articulate enlarged apical nails, inner
ramus of uropod 1 with two accessory nails, outer
ramus of uropod 1 with 8-10 dorsal spines, inner
with 2-3, outer ramus of uropod 2 with 3-6 dorsal
spines, inner with 0-2 dorsomedial spines; peduncle
of uropod 1 with 6-7 apicolateral spines and 3-6
basofacial setae, medially with 6-7 marginal setae
and spines, apicalmost an enlarged spine; peduncle
of uropod 2 with 7-11 dorsal spines, medially with
one apical spine. Peduncle of uropod 3 with 7-10
ventral spines, dorsally with one lateral spine, one
medial spine; rami submasculine, inner extending
to M. 90-110 on article 1 of outer ramus, apex with
2 setae, medial and lateral margins setose, article 2
of outer ramus short, 0.13-0.20, bearing 2 long to
medium setae, medial margin of article 1 setose,
lateral margin with 5 acclivities, spine formula =
2-1-1-1-1-1, or l -1 - l - l - l - i , setal formula = 0-1-
2-1-1-2 or 0-1-1-1-1-2. Telson ordinary, length-
width ratio = 1:1, not fully cleft, each apex wide,
rounded, lateral acclivity shallow, narrow, weak,
NUMBER 245 219
PI
FIGURE 95.?Birubius myallus, new species, holotype, female "a," 9.8 mm (v ?? male "v," 5.7 mm;
w ? female "w," 5.4 mm; y ?? male "y," 7.6 mm).
220 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
bearing short lateral setule, spine next medial
shorter or of length equal to setule, midlateral set-
ules diverse. Cuticle with ordinary bulbar setules.
DESCRIPTION OF MALE.?Apex of rostrum nar-
rower than in female. Eyes enormous, distending
head. Article 2 of antenna 1 with 8 ventral setae;
primary flagellum with 10-13 articles, one calceolus
each on articles 1-7, aesthetascs strongly developed;
accessory flagellum with 8-9 articles. Facial spine
formula on article 4 of antenna 2 = 3-4-5, on arti-
cle 5 = 1-2 or 2-1, article 5 with 4 dorsal sets of
male setae and 3 calceoli, ventrodistal apex with 2
thin spines and one setule; flagellar formula = 28 +
(broken), (1), 3, 4, 6, 8 . . . 28 (broken). Right lacinia
mobilis elaborately sculptured; right rakers 11; left
12 or 13; molars with 6 long and one short spine
apically; basofacial setal formula of article 3 on
mandibular palp = (3-4) + 2 (right) and (2-3) + 2
(left). Palp of maxilla 1 with 6-7 spines and 8-9
setae. Inner plate of maxilla 2 with 2-A medial
setae. Outer plate of maxilliped with only one
apicolateral seta; palp article 2 with 2-3 lateral
setae. Coxa 4 broadened but smaller in relation to
coxa 1 than in female. Facial and setal-spine form-
ulas of pereopods 1-2 on article 4 = 7 and 5, on
article 5 = 6 and 8, on posteroproximal margin of
article 5 = 3-4 and 2-3. Article 2 of pereopods 3-5
as in female but article 5 of pereopod 5 with special
stout posterodistal spine; article 6 narrowed, medial
apex of article 6 on pereopod 5 with (male "y") or
without (male "v") apical digits. Epimera 1-2 broad-
ened; posterior margin of epimeron 3 shortened,
posteroventral tooth absent in male "y" or with
small blunt cusp and indeterminate crenulations in
male "v"; setal formulas, epimeron 1 anteroven-
tral = 9, posteroventral = 4-5, posterior pair set
vertically, epimeron 2 facial = 10-11, epimeron 3
posterior = 3 plus 2 setules and 2 posterodorsal
setules, facial = 3 (or fewer in smaller male), ven-
tral = 4. Articulation line of urosomite 1 distinct.
Spine formula of uropods, uropod 1 peduncle
apicolateral = 5, basofacial = 3, medial = 5 fine
spines and one enlarged spine, uropod 2 peduncle
dorsal = 7-8, dorsal spines on outer ramus of uro-
pod 1 = 6-7, of uropod 2 = 3, inner ramus of
uropod 1 = 1-2, of uropod 2 = one vestigial; ven-
tral spines on peduncle of uropod 3 = 7-8, spine
formula on article 1 of outer ramus = 1-1-1-1-1
2-2-2-2-2, setal formula = 1-1-2-2-1-1-1-1-1-1.
Telson slightly elongate, distal spines not shortened.
OBSERVATIONS.?Right lacinia mobilis of holo-
type apparently gerontic, younger females with
normal lacinia mobilis composed of bifid, shortened
distal branch, and simple, denticulate proximal
branch; holotype with aberrant 2 cones on one lobe
of lower lip; female "u," 8.6 mm, with 11 apico-
lateral spines on peduncle of uropod 1 and with
accessory apical spine on inner ramus of uropod 2.
DESCRIPTION OF JUVFNILE (young females).?Gen-
erally similar to holotype as described but following
spine and seta counts showing lower densities; if
two variations quoted, first based on 5.6 mm female
"n," second based on 5.4 mm female "w": antenna
2 article 4 spine formula = 1-3-4-5, spine formula
of article 5 = 2-2, article 5 with only 3 posterior
setal groups. Right mandible with 7 rakers + one
rudimentary or 9 + 1, molar with 6 spines plus one
disjunct or 5 + 1; left mandible with 8 rakers + 2
rudimentary, molar with 6 spines + one disjunct,
or 5 + 1; article 2 of mandibular palp with 0 or 1
basofacial seta, article 3 of mandibular palp with
basofacial setae in formulas of 1 + 1 or 2 + 1.
Palp of maxilla 1 with only 4 or 5 of main spines.
Outer plate of maxilla 2 with 4 lateral setae, inner
setae like holotype. Outer plate of maxilliped with
3 lateral setae; inner plate and palp article 4 like
holotype but inner plate with only 2 stout apical
spines. Uropod 1 with 4 or 3 basofacial setae, pe-
duncle with 5 or 4 apicolateral spines, outer ramus
with 5 or 6 spines, inner with 2; peduncle of uropod
2 with 8 spines, outer ramus with 4, inner with 1.
Epimeron 1 on female "n" with 5 posterofacial
setae, 3 arranged in vertical row (see illustration),
female "w" with 4 setae, 2 arranged vertically; epi-
meron 2 with 10-11 setae; epimeron 3 widi many
fewer setae than holotype (see illustrations). Right
lacinia mobilis with short distad ramus weakly
bifid (flattened in illustration but generally similar
to unflattened holotype views). Smaller female
therefore advanced in number of spines on outer
ramus of uropod 1 and rakers of left mandible and
basofacial setae on mandibular palps but less ad-
vanced than larger female in other features men-
tioned above.
ILLUSTRATIONS.?Left mandible shown with molar
folded back; one apical seta on article 2 of mandi-
bular palp missing; inner plate of maxilla 1 with
one large apical seta absent, other side bearing this
seta; upper lip viewed slightly anterodorsally; tel-
son of female "w" shown for aberrant condition of
NUMBER 245 221
FIGURE 96.?Birubius myallus, new species, holotype, female "a," 9.8 mm (n = female "n," 5.6
mm; if ? male "v," 5.7 mm; w ? female "w," 5.4 mm; y ? male "y," 7.6 mm).
222 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 97.?Birubius myallus, new species, female "g," 6.8 mm (u - female "u," 8.6 mm).
setation; pereopod 1 of male "y" with facial setae
on article 5 omitted.
HOLOTVPE.?NMV, female "a," 9.8 mm.
TYPE-LOCALITY.?CPBS 10E/2, 5 Apr 1965, West-
ern Port, Victoria, Australia, 3m, fine sand, mud.
VOUCHER MATERIAL (all illustrated).?PPBES
986/5, male "y," 7.6 mm; CPBS 24N/872, male "v,"
5.7 mm; CPBS 25S/4, female "n," 5.6 mm; CPBS
25S/2, female "w," 5.4 mm; PPBES 968/2, female
"u," 8.6 mm; PPBES 968/5, female "g," 6.8 mm.
REMARKS.?Apparently the holotype of this spe-
cies is gerontic, as evidenced by the enlarged eyes,
submasculine uropod 3, supernumerary facial spines
and flagellar articles on antenna 2, supernumerary
setae on mandibular palps and other appendages,
extra cone on one lobe of lower lip, extra (3 instead
of 2) main apical spine on inner plate of maxil-
liped, presence of 2 spines on inner ramus of uro-
pod 2 and reduction in length of spines on telson,
absence of one apical seta on inner plate of maxilla
1 and of the disjunct spine on the right molar, and
the smaller number of other molarial spines and
mandibular rakers.
RELATIONSHIP.?The greater B. myallus-muldar-
pus (4-9) group differs from the B. panamunus-
nammuldus (1-3) group in the absence of long ven-
tral setae on coxa 4. This species and that to follow,
B. kareus (5), form a subgroup in the greater B.
myallus group bearing ventral setae on epimeron 3.
They form a pair of species closely similar to the B.
gambodeni (20) and B. jirrandus (30) groups but
differ from those species in the presence of an ac-
cessory apical nail on the inner ramus of uropod
1, and 2 or more stout apical spines on the inner
plate of the maxilliped. In other respects B. myallus
and B. kareus differ from the B. gambodeni group
in the elongate anterior ridge on pereopod 5 and
the elongate wrists of the gnathopods. From B.
munggai (27) and the B. lowannus (22) group, these
two species differ in the uncrowded setae on epi-
meron 2 and the well setose basofacial peduncle of
uropod 1 and from the B. lowannus group, in addi-
tion, by the long anterior ridge on pereopod 5.
From the B. bate'ii (24) group, these two species
differ in the poorer development of posterior setae
on epimeron 3 and the presence of fully facial setae
on epimeron 3.
MATERIAL.?PPBES, 3 samples from 2 stations
(4); CPBS, 4 samples from 2 stations (5).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 4-9 m, sand, muddy sand.
5. Birubius kareus, new species
FIGURES 98-100
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total body length, greatest width about 60
percent of length; rostrum constricted, narrow,
reaching middle of article 2 on antenna 1. Eyes
NUMBER 245 223
FIGURE 98.?Birubius karexu, new species, holotype, female Ma," 4.6 mm (c ? female "c," 4.2 mm;
m ?m male "m," 5.1 mm; Y ?= left lacinia mobilis).
224 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
mEh
FIGURE 99.?Birubius kareus, new species, holotype, female "a," 4.6 mm (m ? male "m," 5.1 mm).
NUMBER 245 225
medium, largely occluded with pigment. Article 1
on peduncle of antenna 1 about 1.45 times as long
as wide, about twice as wide as article 2, ventral
margin with about 9-10 setules, weakly produced
dorsal apex with 2 setules; article 2 about 0.8 times
as long as article 1, with ventral cycle of 4?6 setae;
primary flagellum with 10-11 articles, about 0.75
times as long as peduncle, bearing short aesthetascs;
accessory flagellum with 7-9 articles. Spine formula
on article 4 of antenna 2 = 1-3-4?3-1 or 1-3-4-3,
dorsal margin with notch bearing 4 setae, ventral
margin with 5-6 groups of 2 long to short setae, one
ventrodistal long spine; article 5 about 0.8 times as
long as article 4, facial spine formula = 1-2-2,
dorsal margin naked, ventral margin with 4 sets of
2 long to short setae, 2 ventrodistal medium spines,
flagellum about 1.0-1.2 times as long as articles 4-5
of peduncle combined, with 10-11 articles. Mandi-
bles with medium palpar hump; right incisor with
3 teeth and accessory cusp; left incisor with 5
humps in 2 branches; right lacinia mobilis bifid,
distal branch shorter than proximal, flabellate,
broad, denticulate, with one facial hump, proximal
branch simple, pointed, with marginal denticles;
left lacinia mobilis with 5 teeth, middle teeth short-
ened; right rakers 8; left rakers 9 plus one rudimen-
tary; molar in form of elongate bulbous hump, right
molar with 5 primarily long spines plus one short
spine strongly disjunct, left molar with 4 primarily
long spines, one short spine strongly disjunct, each
molar with plume; palp article 1 short, article 2 with
one long inner apical seta and 2 other short inner
setae, article 3 about as long as article 2, oblique apex
with 8?10 spine-setae, basofacial formula = 1-1 or
2?1. Inner plate of maxilla 1 ordinary, bearing one
long apical pluseta, one similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
4-5 apicalmedial marginal spines and 5 submarginal
setae. Plates of maxilla 2 extending equally, outer
broader than inner, outer with 4?5 apicolateral setae,
inner with one medial seta. Inner plate of maxilliped
with 2 large, thick apical spines, 3 apicofacial setae,
6-7 medial setae; outer plate with 7-8 medial and
apical spines, 1-2 apicolateral setae; palp article 1
with apicolateral seta, article 2 with 2 apicolateral
setae, medial margin of article 2 moderately setose,
article 3 with 4 facial setae, 2 lateral setae, nail of
article 4 medium, with 2 accessory setules. Coxa 1
strongly expanded distally, anterior margin weakly
concave; main ventral setae of coxae 1^ 4 = 7-7-7-0,
posteriormost seta of coxae 1-2 slightly shortened;
anterior and posterior margins of coxa 4 weakly
divergent, posterior margin almost straight, postero-
dorsal corner sharply rounded, posterdorsal margin
short, concave, width-length ratio of coxa 4 = 7:8.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = (O-l)-(2-3)-3-3, long an-
teriors = 4-5-0-0, short anteriors = 1-4-2-1, no
others. Gnathopods ordinary; width ratios of articles
5-6 on gnathopods 1-2 = 31:37 and 31:39, length
ratios = 77:73 and 69:70; palmar humps ordinary,
palms oblique; article 5 of gnathopods 1-2 elongate,
ovate, posterior margin flat-rounded. Pereopods 1-2
similar; facial setae formula on article 4 = 4 and 4,
on article 5 = 6 and 6; main spine of article 5 extend-
ing to M. 80 on article 6, article 5 with 2 proximo-
posterior spines; spine formula of article 6 = 4 + 4
plus middistal seta, 2 spines especially long; accliv-
ity on inner margin of dactyls of pereopods 1-2
weak, emergent setule long, midfacial pluseta ordi-
nary. Coxae 5-7 posteroventral setule-seta form-
ula = 2-2-1. Articles 4-5 of pereopods 3-4 of ordi-
nary width, facial spine rows moderately developed,
facial ridge formula on article 2 of pereopods
3-5 = 0-2-2, posterior ridge on pereopod 4 short;
width ratios of articles 2, 4, 5, 6 of pereopod 3 =
46:40:34:17, of pereopod 4 = 67:37:23:11, of pe-
reopod 5 = 88:23:21:9, length ratios of pereopod
3 = 76:34:37:43, of pereopod 4 = 89:51:43:51, of
pereopod 5 = 110:28:24:30; article 2 of pereopod
5 reaching middle of article 5; medial apex of
article 6 finely combed and bearing 6 digital proc-
esses. Posteroventral corner of epimeron 1 sharply
protrusive, posterior margin straight, with setule,
anteroventral margin with 6-7 medium setae, pos-
teroventral face with 2-3 long setae; posteroventral
corner of epimeron 2 weakly protuberant, guarded
by setule sinus, posterior margin slightly concave,
facial setae = 7-8; posteroventral corner of epime-
ron 3 with small to medium tooth, with setule sinus,
posterior margin straight, with 1-2 setule notches,
1-2 long setae distally, ventral margin with 2-3
facial setae pointing posteriorly, evenly spread, face
with oblique horizontal row of 4?5 setae and occa-
sionally with another row of 2 setae more ventrally;
epimera 1-3 with small seta on posterodorsal mar-
gin. Urosomite 1 naked, articulation line incom-
plete; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1-2 with articulate enlarged
apical nails, inner ramus of uropod 1 with 2 acces-
226 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 100.?Birubius kareus, new species, holotype, female "a," 4.6 mm (m ? male "m," 5.1
mm; q ? right side converted to left view; y -> female "y," 5.1 mm).
NUMBER 245 227
sory nails, outer ramus of uropod 1 with 6-7 dorsal
spines, inner with 2, outer ramus of uropod 2 with
3-5 dorsal spines, inner with 1-2 dorsomedial
spines; peduncle of uropod 1 with 3-4 apicolateral
spines and 4 basofacial setae, medially with 4-5
marginal setae and spines, apicalmost an enlarged
spine; peduncle of uropod 2 with 7-8 (rarely 12)
dorsal spines, medially with one apical spine. Pe-
duncle of uropod 3 with 6 ventral spines, dorsally
with one lateral spine, one each medial spine and
setule; rami submasculine, inner extending to M. 95
on article 1 of outer ramus, apex with one seta, me-
dial and lateral margins with 2 and 4 setae, article
2 of outer ramus short, 0.21, bearing 2 long setae,
medial margin of article 1 with 3 setae, lateral mar-
gin with 4 acclivities, spine formula = 1-1-1-1-1,
setal formula = 1-1-1-1-2. Telson long, length-
width ratio = 13 : 11, not fully cleft, each apex wide,
rounded, lateral acclivity shallow, weak, bearing
ordinary lateral setule, spine next medial longer
than setule, midlateral setules diverse. Cuticle with
ordinary bulbar setules.
DESCRIPTION OF MALE.?Head shorter and
broader than in female, distended by greatly en-
larged eyes clear of pigment. Article 2 of antenna
1 with 7 ventral setae; primary flagellum with 12
articles, one calceolus each on articles 1?4 or 1-6
(5-6 rudimentary), aesthetascs strongly developed;
accessory flagellum with 8 articles. Facial spine
formula on article 4 of antenna 2 = 1-3-4-3-1 or
1-2-4-3-1, on article 5 = 2-2; article 5 with 3
dorsal sets of male setae and one calceolus, ventro-
distal apex with 2 thin spines and one setule; fla-
gellar formula = 32, 2, 3, 5, 7 . . . 27. Each molar
with 5 spines and one small disjunct spine; baso-
facial setal formula of article 3 on mandibular
palp = 1-2. Inner plate of maxilla 1 very thin,
with 2 large and one small setae. Article 4 of
maxillipedal palp with only one accessory setule.
Coxa 4 broadened but smaller in relation to coxa 1
than in female. Long posterior setae on article 2 of
gnathopods 1-2 = 1-2. Article 2 of only pereopods
4-5 narrower than in female, articles 4?6 of pereo-
pod 5 narrowed, article 5 with falcate posterodistal
spine. Epimera 1-3 broadened; posterior margin of
epimeron 3 slightly shortened; setal formulas, epi-
meron 1 anteroventral = 7, posteroventral = 4, epi-
meron 2 facial = 7, epimeron 3 posterior = 2 and
1-2 setules, facial = 1-2, ventral = 4-5. Spine
formulas of uropods, uropod 1 peduncle apico-
lateral = 3, basofacial = 4, medial = 5-6, uropod 2
peduncle dorsal = 12; dorsal spines on outer ramus
of uropod 1 = 7, of uropod 2 = 5, inner ramus of
uropod 1 = 3, of uropod 2 = 2; ventral spines on
peduncle of uropod 3 = 3; spine formula on article
1 of outer ramus = 1-1-2-2-2-2-2, setal formula =
1-1-1-1-1-1-1. Telson elongate, broadened, distal
spines shortened.
ILLUSTRATIONS.?Female: Upper and lower lips
and maxilla 1 as shown for B. myallus (4), inner
lobes of lower lip swollen perhaps owing to preser-
vational effect; shape of coxae 2-3 as shown for B.
myallus; maxilliped, except for articles 3-4 of palp
and notes in description, as shown for B. panamu-
nus (1).
Male: Urosome in poor condition, not illus-
trated; epimeral view based on partial reconstruc-
tion presented as left view of right epimera; one
seta on inner plate of maxilla 1 hidden.
HOLOTYPE.?NMV, female "a," 4.6 mm.
TYPE-LOCALITY.?CPBS 31N/769, 15 Jul 1969,
Western Port, Victoria, Australia, 15?19 m, fine
sand and mud.
VOUCHER MATERIAL (all illustrated).?CPBS 32S/
866, male "m," 5.1 mm; CPBS 21S/4, female "c,"
4.2 mm; CPBS 31N/867, female "y," 5.1 mm.
RELATIONSHIP.?This species and Birubius myal-
lus (4) are extremely close morphologically; the
best distinction is the wide degree of disjunction of
the clear spine on the mandibular molars which in
B. myallus is closely approximate to the other
spines but which in B. kareus is widely disjunct; the
male of B. kareus bears a peculiar falcate spine on
article 5 of pereopod 5 which in males of B. myallus
is smaller and scarcely modified. Subtle differences
in the shape of the head from dorsal view are ap-
parent between the species, B. kareus having a
slightly more constricted and streamlined rostrum
with the eyes of the male fully contiguous dorsally
whereas the eyes of male B. myallus, though ex-
tremely large, do not meet dorsally and are ex-
tended into nacelles laterodorsally. Birubius kareus
has slightly stouter gnathopods than those of B.
myallus and the distal branch of the right lacinia
mobilis is multifid whereas in B. myallus that
ramus is at best bifid or weakly so. All specimens of
B. myallus except female "n," 5.6 mm, have at least
one basofacial seta on article 2(!) of the mandibular
palp whereas these setae are absent in B. kareus.
These distinctions are extremely weak and make the
228 SMITHSONIAN CONTRIBUTION'S TO ZOOLOGY
routine identification of these two species very diffi-
cult. We trust that future collections of these ani-
mals may clarify any further distinctions or demon-
strate that they are but phenotypes of a single
species.
MATERIAL.?CPBS, 35 samples from 13 stations
(54); WPBES, 2 samples from 2 stations (2).
DISTRIBUTION.?Victoria, Western Port, 9-19 m,
sand, silty sand.
6. Birubius apart, new species
FICURES 101-104
DESCRIPTION OF FEMALE.?Head about 17 percent
of total body length, greatest width about 72 per-
cent of length; rostrum unconstricted, broad, reach-
ing middle of article 2 on antenna 1. Eyes medium,
largely occluded with pigment. Article 1 of pedun-
cle on antenna 1 about 1.4 times as long as wide,
about twice as wide as article 2, ventral margin
with about 7 setules, weakly produced dorsal apex
with 4 setules; article 2 about 0.72 times as long as
article 1, with ventral cycle of 4 setae; primary fla-
gellum with 10 articles, about as long as peduncle,
bearing long aesthetascs; accessory flagellum with
8 articles. Spine formula of article 4 on antenna
2 = 1-3-4-3, dorsal margin bearing 2 setae, ventral
margin with 4 groups of 1-2 long to medium setae,
one ventrodistal long spine; article 5 about 0.71
times as long as article 4, facial spine formula = 2,
dorsal margin naked, ventral margin with 2 sets of
one long seta and one setule, 3 ventrodistal long to
medium spines and setae; flagellum about 1.4 times
as long as articles 4-5 of peduncle combined, with
11-12 articles. Mandibles with weak palpar hump;
right incisor with 3 teeth and notch; left incisor
with 4 humps in 2 branches; right lacinia mobilis
bifid, distal branch little shorter than proximal, fla-
bellate, broad, subbifid or denticulate, with one
facial hump, proximal branch simple, pointed, widi
marginal denticles; left lacinia mobilis with 4 teeth;
right rakers 6 plus one rudimentary; left rakers 7;
molars bulbous, each with 5 primarily long spines
plus one short spine not disjunct; palp article 1
short, article 2 with one long inner apical seta and
2 other short inner setae, article 3 about 1.1 times
as long as article 2, oblique apex with 9 spine-
setae, basofacial formula = 0-1. Inner plate of
maxilla 1 especially thin, bearing one long apical
pluseta, one similar apicomedial seta, 2 apicolateral
much shorter setae; palp article 2 with 5 apical-
medial marginal spines and 5 submarginal setae.
Plates of maxilla 2 extending equally, outer scarcely
broader than inner, outer with 3 apicolateral setae,
inner with one medial seta. Inner plate of maxil-
liped with 3 large, thick apical spines, 4 apicofacial
setae, 5 medial setae; outer plate with 8 medial and
apical spines, one apicolateral seta; palp article 1
with apicolateral seta, article 2 with 3 groups of one
each apicolateral setae, medial margin of article 2
moderately setose, article 3 scarcely protuberant,
with 4 facial setae, one lateral seta, nail of article 4
medium, with 2 accessory setules. Coxa 1 scarcely
expanded distally, anterior margin weakly convex;
main ventral setae of coxae 1-4 = 4-4-(3-4)-0,
posteriormost seta of roxae 1-3 slightly shortened;
anterior and posterior margins of coxa 4 slightly
divergent, posterior margin almost straight, postero-
dorsal corner rounded, posterodorsal margin me-
dium, concave, undulant, width-length ratio of
coxa 4 = 4:5. Long posterior setae of gnathopods
1-2 and pereopods 1-2 = 1-3-3-4, short anteriors =
1?1?(3-5>-l, long anteriors = (5-6)-7-0-0. short
posteriors = 1-0-1-2. Gnathopods ordinary, width
ratios on articles 5-6 of gnathopods 1-2 = 25:34
and 28:34, length ratios = 78:58 and 78:59; pal-
mar humps ordinary, palms weakly oblique; article 5
of gnathopod 1 elongate, ovate, posterior margin flat,
article 5 of gnathopod 2 elongate, ovate, posterior
margin rounded. Pereopod 2 similar but stouter
than pereopod 1, especially on article 4, facial setae
formula on article 4 = 4 and 3, on article 5 = 4 and 5;
main spine of article 5 extending to M. 80 on arti-
cle 6, article 5 lacking proximoposterior spines; spine
formula of article 6 = 3 + 4 and 4 + 4 plus mid-
distal seta, some spines especially long; acclivity on
inner margin of dactyls of pereopods 1-2 sharp,
produced as tooth, emergent setule long, midfacial
pluseta ordinary. Coxae 5?7 posteroventral setule
formula = 1-1-1. Articles 4-5 of pereopods 3?4
broad to ordinary, facial spine rows sparse to mod-
erately developed, facial ridge formulas of article 2
on pereopods 3-5 ? 0-2-2, width ratios of articles
2, 4, 5, 6 of pereopod 3 = 44:47:41:19, of pereo-
pod 4 = 73:42:28:13, of pereopod 5 = 88:20:16:9,
length ratios of pereopod 3 = 81:36:36:40, of
pereopod 4 = 86:57:40:49, of pereopod 5 = 112:25:
22:20; article 2 of pereopod 5 reaching middle of
article 5; medial apex of article 6 bearing 4 digital
NUMBER 245 229
FIGURE 101.?Birubius apart, new species, male "a," 3.9 mm.
processes. Posteroventral corner of epimeron 1
rounded, posterior margin convex, anteroventral
margin with 3-4 medium setae, posteroventral face
with 3 long setae, posterior pair set vertically;
posteroventral corner of epimeron 2 rounded, pos-
terior margin convex, facial setae = 6-7; postero-
ventral corner of epimeron 3 rounded, with small to
medium tooth, bearing setule, posterior margin
straight, ventral margin naked, face with oblique
row of 6-8 thick setae commencing on tooth; epimera
1-3 with setule on posterodosal margin. Urosomite 1
with one lateral and one ventral setule at base of
uropod 1, articulation line incomplete; urosomite 3
weakly protuberant dorsally. Rami of uropods 1-2
230 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 102.?Birubius apart, new species, male "a," S.9 mo).
NUMBER 245 231
with articulate enlarged apical nails, inner ramus of
uropod 1 with two accessory nails, outer ramus of
uropod 1 with 3-4 dorsal spines, inner with one,
outer ramus of uropod 2 with 1-2 dorsal spines,
inner with one dorsomedial spine; peduncle of uro-
pod 1 with 2 apicolateral spines, and one tiny baso-
facial setule, medially with 2 marginal spines,
apicalmost enlarged; peduncle of uropod 2 with 4-5
dorsal spines, medially with one small apical spine.
Peduncle of uropod 3 with 5 ventral spines, dorsally
with one lateral spine, one medial spine; rami
feminine, inner extending to M. 85 on article 1 of
outer ramus, apex with one seta, medial and lateral
margins naked, article 2 of outer ramus ordinary,
0.22, bearing 2 long setae, apicomedial margin of
article 1 naked, lateral margin with 2 acclivities,
spine formula = 1-1-1, setal formula = 1-1-1. Tel-
son long, length-width ratio = 9:7, not fully cleft,
each apex of medium width, rounded, lateral accli-
vity broad, shallow, bearing ordinary lateral setule,
spine next medial shorter than setule, midlateral
setules diverse. Cuticle with ordinary bulbar setules.
OBSERVATIONS (female).?Prebuccal complex con-
vex anteriorly, poorly extended, epistome and
upper lip partially fused but weak dorsal articula-
tion line present, anterior face of upper lip with
weak rugosity marked as crescent from anterior
view, ventral margin weakly excavate. Holotype
immature, bearing rudimentary brood plates only
on pereonite 4.
ILLUSTRATIONS (female).?Following parts similar
to those of male: prebuccal, upper lip and epistome,
mandibular palp, left mandible (except basofacial
formula), molars, lower lip, maxillae 1-2, maxilli-
ped, gnathopods 1-2, pereopods 3-4.
DESCRIPTION OF MALE.?Head about 20 percent
of total body length, greatest width about 65 per-
cent of length; rostrum weakly constricted, broad,
reaching middle of article 2 on antenna 1. Eyes
medium, clear of occluding pigment, eosin in color.
Article 1 of peduncle on antenna 1 about 1.25 times
as long as wide, about 3 times as wide as article 2,
ventral margin with about 16 setules, produced
dorsal apex with 6 setules, medial face with patch
of fuzz; article 2 about 0.8 times as long as article
1, with ventral cycle of 4 setae; primary flagellum
with 8-10 articles, about 0.85 times as long as pe-
duncle, bearing calceoli on articles 1-5; accessory
flagellum with 6-7 articles. Spine formula of article
4 on antenna 2 = 3-^-3, dorsomedial margin of
articles 3-4 bearing fuzz, ventral margin with 4
groups of 1-2 short to medium setae, one ventro-
distal long spine; article 5 about as long as article 4,
facial spine formula = 2, dorsal margin bearing 3
sets of male setae and 2 calceoli, ventral margin
with 2 sets of 1-3 short to medium setae, 3 ventro-
distal short setae; flagellum elongate, flagellar form-
ula = 28, 2, 3, 5, 7 . . . 27. Mandibles with medium
palpar hump; right incisor with 3 teeth, accessory
hump, 2 notches; left incisor with 5 humps in 2
branches; right lacinia mobilis bifid, distal branch
little shorter than proximal, flabellate, subbifid,
denticulate, proximal branch simple, pointed, with
marginal denticles and facial humps; left lacinia mo-
bilis with 4 very low teeth; right and left rakers 6
plus 3 rudimentaries; molars composed of bulbous
humps, right molar with 7 primarily medium
spines, one short spine weakly disjunct, left molar
with 6 primarily medium spines, one short spine
weakly disjunct; palp article I short, article 2 with
one long inner apical seta and 2 other short inner
setae, article 3 about 1.15 times as long as article 2,
oblique apex with 10 spine-setae, basofacial form-
ula = 2-2. Inner plate of maxilla 1 especially thin,
bearing one long apical pluseta, one similar apico-
medial seta, 2 apicolateral much shorter setae; palp
article 2 with 5 apicalmedial marginal spines and 5
marginal setae. Plates of maxilla 2 extending
equally, subequally broad, outer with 3 apicolateral
setae, inner with one medial seta. Inner plate of
maxilliped with 2 large thick, apical spines, 2 apico-
facial setae, 4 medial setae; outer plate with 6 me-
dial and apical spines, one apicolateral seta; palp
article 1 with apicolateral seta, article 2 with 3
groups of 1-2 apicolateral setae, medial margin of
article 2 strongly setose, article 3 weakly protu-
berant, with 4 facial setae, 2 lateral setae, nail of
article 4 long, with 2 accessory setules. Coxa 1
scarcely expanded distally, anterior margin weakly
concave; main ventral setae of coxae 1-4 = 4?(3?4)-
3-0, posteriormost seta of coxae 1-2 slightly short-
ened; anterior and posterior margins of coxa 4
almost divergent, posterior margin oblique, almost
straight, posterodorsal corner rounded, posterodorsal
margin short, ordinary, width-length ratio of coxa
4 = 5:6; long posterior setae of gnathopods 1-2 and
pereopods 1-2 = 1-2-2-3, short anteriors = 1-2-2-3,
short anteriors = 1?1?4?2, short posteriors = 1-0-
1-3. Gnathopods thin, almost identical; width ratios
on articles 5-6 of gnathopods 1-2 = 29:32 and
232 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 103.?Birubius apart, new species, male "a," 3.9 mm (J ? male "f," 3.8 mm).
28:33, length ratios = 80:60 and 84:60; palmar
humps ordinary, palms weakly oblique; article 5 of
gnathopods 1-2 elongate, ovate, posterior margin
flat, long. Pereopods 1-2 similar; facial setae form-
ula on article 4 = 4 and 3, on article 5 = 5 and 5;
main spine of article 5 extending to M. 100 on arti-
cle 6, article 5 lacking proximoposterior spines;
spine formula of article 6 = 3 + 4 plus middistal
seta, some spines especially long; acclivity on inner
margin of dactyls of pereopods 1-2 weak, emergent
setule long, midfacial pluseta ordinary. Coxae 5-7
posteroventral setule formula = 1-1-0. Articles 4-5
of pereopods 3-4 of ordinary width, facial spine
rows sparse, facial ridge formulas of article 2 on
pereopods 3-5 = 0-2-2, width ratios of articles
2, 4, 5, 6 of pereopod 3 = 40:39:33:17, of pereopod
4 = 63:38:22:9, of pereopod 5 = 83:17:14:6, length
ratios of pereopod 3 = 78:34:42:35, of pereopod
4 = 89:58:42:50, of pereopod 5 = 123:27:23:20;
article 2 of pereopod 5 reaching middle of arti-
cle 5, latter with 2 special posteroventral spines;
medial apex of article 6 weakly scalloped.
Posteroventral corner of epimeron 1 rounded, pos-
terior margin convex, corner with setule, antero-
ventral margin with 4-5 medium setae, postero-
ventral face with 3-5 medium setae, in tandem or
partially paired vertically; posteroventral corner of
epimeron 2 rounded, posterior margin convex,
facial setae = 5-8 crowded anteriorly in terminal
adult; posteroventral corner of epimeron 3 with
setule sinus, small to medium tooth, posterior mar-
gin weakly convex, face with oblique row of 8-11
setae near posterior margin, setae divided into
groups of 5-3-2 or 5-2-1 or 6-5 or 5-2-2 (or other
variations); epimera 1-3 with setule on postero-
dorsal margin. Urosomite 1 naked, articulation line
complete; urosomite 3 strongly protuberant dor-
sally. Rami of uropods 1-2 with articulate, en-
larged apical nails, inner ramus of uropod 1 with
two accessory nails, outer ramus of uropod 1 with
3 dorsal spines, inner with one, outer ramus of
uropod 2 with 3 dorsal spines, inner with one
dorsomedial spine; peduncle of uropod 1 with 2
apicolateral spines and one basofacial seta, me-
dially with 4 marginal setae and spines, apicalmost
an enlarged spine; peduncle of uropod 2 with
NUMBER 245 233
FIGURE 104.?Birubius apart, new species, holotype, female "a," 3.75 mm (j = juvenile "j,"
3.46 mm).
234 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
10-11 dorsal spines, medially with one small apical
setule. Peduncle of uropod 5 with 6 ventral spines,
dorsally with one lateral spine, one medial spinule;
rami masculine, inner extending to M. 100 on arti-
cle 1 of outer ramus, apex with 2 setae, medial and
lateral margins setose, article 2 of outer ramus
ordinary, 0.16, bearing 2 long setae, medial margin
of article 1 setose, lateral margin with 5 acclivities,
spine formula = 0-1-1-1-1-2, setal formula = 1-1-
1-1-1-1. Telson especially long, length-width
ratio = 8:5, not fully cleft, each apex narrow,
rounded, lateral acclivity deep, bearing ordinary
lateral setule, spine next medial of length equal to
setule, midlateral setules diverse, each lobe with
dorsal row of denticles. Cuticle with sparse ordi-
nary bulbar setules.
OBSERVATIONS (male).?Male "m" with 8 ventral
setae on epimeron 2.
ILLUSTRATIONS (male).?Eyes shrunken ventrad so
dorsal margin from lateral view appearing flat;
antenna 1 constructed of flagella from right and left
members fitted onto left peduncle; left mandibular
molar like right.
VARIATIONS.?Calceoli often lost through damage
or occasionally absent, for example, absent on right
antenna 1 of male "a".
HOLOTYPE.?AM, female "a," 3.75 mm.
TYPE-LOCALITY.?EBS 31, 25 Apr 1972, Jervis
Bay, Murray's Beach, New South Wales, Australia,
6 m, Halophila.
VOUCHER MATERIAL.?EBS 20: juvenile ")," 3.46
mm (illus.); juvenile "c," 3.21 mm. EBS 22: juve-
nile "d," 3.95 mm; juvenile "e," 3.51 mm. AM
P.18125: male "a," 3.9 mm (illus.); male "f," 3.8
mm (illus.). PPBES 1834: male "m," 4.45 mm.
PPBES 1842: female "n," 4.30 mm. PPBES 1811:
subadult male "t," 4.05 mm.
RELATIONSHIP.?This species is distinguished
from the B. myallus (4) subsection in the absence
of ventral setae on epimeron 3 and the reduction
of basofacial setae on uropod 1.
Several discrepancies in morphology suggest that
the two sexes are distinct species: the unconstricted
rostrum of the female, the much narrower coxa 4,
the shorter main spine on article 5 of pereopods
1-2, the sharp inner tooth on the dactyls of pereo-
pods 1-2, and the shorter article 2 of antenna 1.
Other differences in shapes and spine counts might
be attributed to size or sexual differences. However,
both sexes have been found together in Victorian
samples after protracted search.
Birubius apari differs from B. panamunus (1)
in the absence of long setae on coxa 4, the absence
of long basofacial setae on uropod 1 and the ab-
sence of a midridge on pereopod 3.
MATERIAL.?EBS, 4 samples from one station (6);
AM, one sample (2); PPBES, 3 samples from 3 sta-
tions (3).
DISTRIBUTION.?New South Wales: Jervis Bay,
Murray's Beach, 6 m, Halophila and Zostera. Vic-
toria: Port Phillip Bay, 0.5-3.5 m, sand and shelly
sand. South Australia: Kangaroo Island, neritic.
7. Birubius car too, new species
Ficutcs 10.V-I07
DESCRIPTION OF FEMALE.?Head about 15 per-
cent of total body length, greatest width about 70
per cent of length; rostrum unconstricted, broad,
reaching middle of article 2 on antenna I. Eyes
medium-large, largely occluded with pigment. Arti-
cle 1 on peduncle of antenna 1 about 1.2 times
as long as wide, about 1.4 times as wide as article
2, ventral margin with about 7 setules, weakly pro-
duced dorsal apex with 4 setules, article 2 about
0.8 times as long as article 1, with ventral crescent
of 9 setae; primary flagellum with 7-8 articles,
about 0.65 times as long as peduncle, bearing long
aesthetascs; accessory flagellum with 5?7 articles.
Spine formula on article 4 of antenna 2 = 1-3-4-4
or 1-3-4-3, dorsal margin with notch bearing 3
setae, ventral margin with 5-6 groups of 1-3 long
to short setae, one ventrodistal long spine; article
5 about 0.75 times as long as article 4, facial spine
formula = 0-2 or 2-1 or 2-2-1, dorsal margin
naked, ventral margin with 4 sets of 1-2 long to
short setae, 3 ventrodistal long to medium spines
and setae; flagellum as long as articles 4?5 of pe-
duncle combined, with 9-11 articles. Mandibles
with weak palpar hump; right incisor with 3 teeth,
occasional notch; left incisor with 4 humps in 2
branches, right lacinia mobilis bifid, distal branch
shorter than proximal, broad, subbifid, proximal
branch simple, pointed, with marginal denticles;
left lacinia mobilis with 4 teeth plus occasional
accessory tooth, middle teeth shortened; right rak-
ers 5-8; left rakers 7-9; molar in form of short
protrusion demarcated mainly by spines, right mo-
lar with 5-7 primarily long spines, none disjunct,
NUMBER 245 235
Gl
FIGURE 105.?Birubius cartoo, new species, holotype, female "a," 4.6 mm.
236 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 106.?Birubius cartoo, new species, holotype, female "a," 4.6 mm.
NUMBER 245 237
left molar with 6-8 primarily long spines, none
disjunct; palp article 1 short, article 2 with one
long inner apical seta and 2-4 other short inner
setae, article 3 about 1.3 times as long as article 2,
oblique apex with 9 spine-setae, basofacial for-
mula = 0-1 or 1-1. Inner plate of maxilla 1 thin,
bearing 2 long apical plusetae, 2 apicolateral much
shorter setae; palp article 2 with 5 apicalmedial
marginal spines and 5 submarginal setae. Inner
plate of maxilla 2 slightly shorter and much nar-
rower than outer, outer with 3 apicolateral setae,
inner with one medial seta. Inner plate of maxil-
liped with 2 large thick apical spines, one ventral
seta, 3 apicofacial setae, 5 medial setae; outer plate
with 6-7 medial and apical spines, no apicolateral
setae; palp article I with apicolateral seta, article 2
with 4 apicolateral setae, medial margin of article
2 moderately setose, article 3 unprotuberant, with
4 facial setae, 1-2 lateral setae, nail of article 4
medium, with 1-2 accessory setules. Coxa 1 not
expanded distally, anterior margin weakly convex;
main ventral setae of coxae 1-4 = 15?(13?14)?10?0,
posteriormost seta of coxae 1-2 shortened; anterior
and posterior margins of coxa 4 almost parallel,
posterior margin almost straight, posterodorsal cor-
ner subsharp, posterodorsal margin ordinary, un-
dulant, width-length ratio of coxa 4 = 7:10. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = (l-3)-(3-4)-5-6, long anteriors =
(4-5)-(5-6)-0-0, others not counted. Gnathopods
thin; width ratios of articles 5?6 on gnathopods
1-2 = 24:29 and 29:29, length ratios = 77:54
and 71:53; palmar humps small, palms almost
transverse; article 5 of gnathopods 1-2 elongate,
trapezoidal, posterior margin flat, long. Pereopods
1-2 similar; facial setae formula on article 4 = 6-6,
on article 5 = 5-5; main spine of article 5 extend-
ing to M. 90 on article 6, article 5 lacking proximo-
posterior spines; spine formula of article 6 = 4 + 6
or 4 -I- 5 or 5 + 6 plus middistal seta, some spines
especially long; acclivity on inner margin of dactyls
of pereopods 1-2 weak, emergent setule long, mid-
facial pluseta ordinary. Coxae 5-7 posteroventral set-
ule formula = 2-l-(0-4). Articles 4-5 of pereopods
3-4 broad, facial spine rows dense, facial ridge for-
mula on article 2 of pereopods 3-5 = 1-2-2, width
ratios of articles 2, 4, 5, 6, of pereopod 3 = 48:48:
45:24, of pereopod 4 = 65:43:27:11, of pereopod 5 =
68:20:16:9, length ratios of pereopod 3 = 82:45:
44:43, of pereopod 4 = 83:55:41:62, of pereo-
pod 5 = 100:24:20:19; article 2 of pereopod 5
almost reaching apex of article 4; medial apex of
article 6 finely combed, bearing 6 digital processes.
Posteroventral corner of epimeron 1 subquadrate,
posterior margin weakly convex, with 2 setules,
anteroventral margin with 9-3 long setae, postero-
ventral face with 2 medium setae; posteroventral
corner of epimeron 2 quadrate, posterior margin
strongly convex, with 2 setules, facial setae = 9-3,
several posterior pairs set obliquely; posteroventral
corner of epimeron 3 with large tooth, posterior
margin almost straight, with 4-6 setules, ventral
margin with 3-6 mainly posterior setae evenly
spread; epimeron 3 with rugosities at posterodorsal
invagination. Urosomite 1 elongate, naked, articu-
lation line complete, forming lateral ridge running
forward to rounded socket fitting invagination of
pleonite 3; urosomite 2 largely hidden but elongate
and marked by dorsal cleft; urosomite 3 weakly
protuberant dorsally. Rami of uropods 1-2 with
articulate enlarged apical nails, inner ramus of
uropod 1 with two accessory nails, one of these
disjunct, outer ramus of uropod 1 with 3-4 dorsal
spines, inner with one, outer ramus of uropod 2
with 1-2 dorsal spines, inner with 0-1 dorsomedial
spine; peduncle of uropod 1 with 2 apicolateral
spines and 2 basofacial setae, medially with 2-3
marginal setae and spines, apicalmost an enlarged
spine; peduncle of uropod 2 with 4?5 (rarely 6)
dorsal spines, medially with one apical spine. Pe-
duncle of uropod 3 with 6-7 ventral spines, dor-
sally with one lateral spine, one medial spine and
setule; rami submasculine, inner extending to M.
90+ on article 1 of outer ramus, apex with 2 setae,
lateral margin setose, medial margin with one seta,
article 2 of outer ramus short, 0.16, bearing 2 long
setae, medial margin of article 1 with 4 setae, lat-
eral margin with 4 acclivities, spine formula =
0-0-0-1-1, setal formula = 2-2-2-1-1. Telson long,
length-width ratio = 7:5, no fully cleft, each apex
of medium width, rounded, lateral acclivity nar-
row, weak, bearing ordinary lateral setule, spine
next medial longer than setule, midlateral setules
highly diverse. Cuticle with closely packed bulbar
setules partially surrounded by alate plaques form-
ing clear spaces, emergent setules especially short.
DESCRIPTION OF MALE.?Apex of rostrum nar-
rowed. Eyes enlarged. Primary flagellum of an-
tenna 1 with 8 articles, bearing calceoli and elon-
gate aesthetascs. Spine formula on article 4 of an-
238 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 107.?Birubius cartoo, new species, holotype, female "a," 4.6 mm (b ? male "b." 4.2 mm;
e = female "c," 4.2 mm; g ? male "g," 3.9 mm; i ? male "i," 3.8 mm; / ? male "j," 3.8 mm;
k ? male "k," 3.9 mm; n = female "n," 4.2 mm; u ?= juvenile "u," $3 mm).
tenna 2 = 3-4-3 or 3-4-4, of article 5 = 2, article
5 with up to 4 dorsal calceoli and 4 clusters of
male setae; flagellar formula = 28, 1, 2, 4, 5, 7,
9 . . . 27 or 1-3, 5,7 . . . . or 2, 3,5,7 . . . . Distal branch
of right lacinia mobilis almost truncate; basofacial
setae of article 3 on mandibular palp = 2 + 1 . Article
6 of pereopod 4 more attenuate, length-width ratio
= 18:3 (female = 16:3); article 5 also more attenu-
ate. Article 2 of pereopods 3-5 not narrower than
in female. Epimera 1-3 broadened; posterior mar-
gin of epimeron 3 not shortened; setal formulas,
epimeron 1 anteroventral = 8, posteroventral =
2-3, epimeron 2 facial = 10, epimeron 3 poste-
rior = 6, tooth shorter, ventral = 4; spine formulas
of uropods, uropod 1 peduncle apicolateral = 2-3,
basofacial = 2, uropod 2 peduncle dorsal = 9, dor-
NUMBER 245 239
sal spines on outer ramus of uropod 1 = 3-4, of
uropod 2 = 2, inner ramus of uropod 1 = 1, of uro-
pod 2 = 1 . Ventral spines on peduncle of uropod
3 = 6; spine formula on article 1 of outer ramus =
1-1-1-1-1-1-1, setal formula = 1-1-1-1-1-1-1.
Telson slightly elongate, distal spines scarcely
shortened.
OBSERVATIONS.?Shape of coxa 7 variable, not
sex-linked, varying from blunt triangle to a more
anteriorly produced form, posterior setae varying
from 0 to 4; six coxae illustrated to show varia-
tions. Inner ramus of uropod 2 in female "c" and
juvenile *'u" with small additional spine, see illus-
tration.
HOIOTYPE.?NMV, female "a," 4.6 mm.
TYPE-LOCALITY.?CPBS 22S/4, 9 Mar 1965,
Western Port, Victoria, Australia, 10.5 m, sand
with mud, shell, stones.
VOUCHER MATERIAL (all illustrated).?PPBES
906/5: male "b," 4.2 mm; female "c," 4.2 mm.
PPBES 986/4: male "g," 3.9 mm. PPBES 919/3:
male "i," 3.8 mm. PPBES 902/4: male " j , " 3.8 mm.
PPBES 127: male "k," 3.9 mm. PPBES 902/3:
female "n," 4.2 mm. PPBES 912/4: juvenile "u,"
3.3 mm.
RELATIONSHIP.?Birubius cartoo, B. thalmus (8)
and B. muldarpus (9) form a group differing from
the B. myallus (4) to B. apari (6) group in the ab-
sence of fully facial setae on epimeron 3, but epi-
meron 3 bears ventral setae unlike B. apari (6).
Birubius cartoo and B. thalmus are very close to
B. panamunus (1) but differ in the absence of setae
on coxa 4 and the absence of fully facial setae on
epimeron 3. Birubius cartoo is unusual in bearing
a midfacial ridge on pereopod 3.
The broadened rostra of the B. cartoo group
suggest that it may be near the ancestral stock of
B. muldarpus (9) and B. gallangus (10), both of
which, however, have reduced tooth and setae of
epimeron 3. See comments written with B. thalmus
(8).
MATERIAL.?CPBS, one sample from one station
(1); PPBES, 46 samples from 18 stations (192).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 2-18 m, sand, silty sand, sandy silt.
8. Birubius thalmus, new species
FIGURES 108, 109
DESCRIPTION OF FEMALE.?Head about 21 percent
of total body length, greatest width about 70 percent
of length; rostrum scarcely constricted, almost reach-
ing middle of article 2 on antenna 1. Eyes medium-
large, largely occluded with pigment. Article 1 on
peduncle of antenna 1 as long as wide, about twice
as wide as article 2, ventral margin with about 7
setules, weakly produced dorsal apex with 3 set-
ules; article 2 as long as article 1, with ventral
crescent of 8 setae; primary flagellum' with 8 arti-
cles, about 0.73 times as long as peduncle, bearing
long aesthetascs; accessory flagellum with 7 articles.
Spine formula on article 4 of antenna 2 = 1-3-4-5,
dorsal margin with notch bearing 3 setae, ventral
margin with 4 groups of 1-2 long to medium setae,
one ventrodistal long spine; article 5 about 0.8
times as long as article 4, facial spine formula = 2,
dorsal margin naked, ventral margin with 3 sets
of 1-2 long to medium setae, 3 ventrodistal long
to medium spines and setae; flagellum about as
long as articles 4-5 of peduncle combined, with 11
articles. Mandibles with medium palpar hump;
right incisor with 3 teeth and notch; left incisor
with 3 humps in 2 branches; right lacinia mobilis
bifid, distal branch shorter than proximal, flabel-
late, denticulate, proximal branch simple, pointed,
with marginal denticles; left lacinia mobilis with
4 teeth, middle teeth shortened; right rakers 8;
left rakers 9; molar in form of bulbous hump, right
molar with 6 primarily long spines, left molar with
7 primarily long spines, none disjunct; palp article
1 short, article 2 with one short inner apical seta
and 3 other short inner setae, article 3 about 1.15
times as long as article 2, oblique apex with 8
spine-setae, basofacial formula = 1-1 or 0-1. Inner
plate of maxilla 1 somewhat thin, bearing one long
apical pluseta, one similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
4 apicalmedial marginal spines and 4 submarginal
setae. Inner plate of maxilla 2 slightly shorter and
narrower than outer, outer with 3 apicolateral
setae, inner without medial setae. Inner plate of
maxilliped with 2 large thick apical spines, 2 apico-
facial setae, 5 medial setae; outer plate with 7 me-
dial and apical spines, no apicolateral setae; palp
article 1 with apicolateral seta, article 2 with 4-5
groups of 1-2 apicolateral setae, medial margin of
article' 2 strongly to moderately setose, article 3
protuberant, with 5 facial setae, 4 lateral setae, nail
of article 4 medium, with 2 accessory setules. Coxa
1 scarcely expanded distally, anterior margin con-
240 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 108.?Birubius thalmus, new species, holotype, female "a," 4.3 mm.
vex; main ventral setae of coxae 1-4 = 8-8-7-0,
posteriormost seta of coxae 1-2 shortened; anterior
and posterior margins of coxa 4 almost parallel,
posterior margin almost straight, posterodorsal cor-
ner rounded, posterodorsal margin short, concave,
width-length ratio of coxa 4 = 5:7. Long poste-
rior setae on article 2 of gnathopods 1-2 and pe-
reopods 1-2 = 4-3~(6-7)-(6-7), short anteriors =
(3-4)-l-3-2, long anteriors = 4-4-0-0. Gnathopods
moderately thin, almost identical; width ratios of
articles 5-6 on gnathopods 1-2 = 26:31 and 26:31,
length ratios = 62:52 and 60:54; palmar humps
small, palms almost transverse; article 5 of gnatho-
pods 1-2 elongate, ovate, trapezoidal, posterior
margin rounded-flat, long. Pereopods 1?2 similar;
facial setae formula on article 4 = 5-5, on article
5 = 5-6; main spine of article 5 extending to M.
100 on article 6, article 5 lacking proximoposterior
spines; spine formula of article 6 = 4 + 5 plus mid-
distal spine, some spines especially long; acclivity
on inner margin of dactyls of pereopods 1-2 weak,
emergent setule long, midfacial pluseta ordinary.
Coxae 5-7 posteroventral setule formula = 3-2-1.
Articles 4-5 of pereopods 3-4 broad, facial spine
rows dense, facial ridge formula on article 2 of
pereopods 3-5 = 1-2-2; width ratios of articles 2,
4, 5, 6 of pereopod 3 = 47:49:45:25, of pereopod
4 = 69:44:25:13, of pereopod 5 = 68:20:16:8, length
NUMBER 245 241
Pi
FIGURE 109.?Birubius thalmus, new species, holotype, female "a," 4.3 mm.
242 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ratios of pereopod 3 = 80:42:44:35, of pereopod
4 = 86:54:36:51, of pereopod 5 = 95:24:22:23;
article 2 of pereopod 5 reaching apex of article
4; medial apex of article 6 finely combed, bear-
ing 6 digital processes. Posteroventral corner of
epimeron 1 subquadrate, posterior margin convex,
with setule, anteroventral margin with 9 long to
medium setae, posteroventral face with 2 long
setae; posteroventral corner of epimeron 2 weakly
protuberant, posterior margin convex, with setule,
facial setae = 8; posteroventral corner of epimeron
3 with small to medium tooth, posterior margin
sinuous, with 4-5 setules, ventral margin with 7
setae evenly spread, pointing posteriorly and dor-
sally, with rugosities at posterodorsal invagination.
Urosomite 1 naked, articulation line complete,
forming lateral ridge running foward to rounded
socket fitting invagination of pleonite 3; urosomite
2 largely hidden but elongate and marked by dor-
sal cleft; urosomite 3 protuberant dorsally. Rami
of uropods 1-2 with articulate enlarged apical
nails, inner ramus of uropod 1 with one accessory
nail, outer ramus of uropod 1 with 3 dorsal spines,
inner with one, outer ramus of uropod 2 with one
dorsal spine, inner with one dorsomedial spine;
peduncle of uropod 1 with 2 apicolateral spines
and 3 basofacial setae, medially with one marginal
seta and apical enlarged spine; peduncle of uropod
2 with 5 dorsal spines, medially with one small api-
cal spine. Peduncle of uropod 3 with 5 ventral
spines, dorsally with one lateral spine, one medial
spine and setule; rami masculine, inner extending
to M. 100 on article 1 of outer ramus, apex with
2 setae, medial and lateral margins setose, article
2 of outer ramus short, 0.15, bearing 2 long setae,
medial margin of article 1 setose, lateral margin
with 4 acclivities, spine formula = 1-1-1-1-1, setal
formula = 0-1-1-1-1. Telson long, length-width
ratio = 17:13, not fully deft, each apex wide,
rounded, lateral acclivity deep, bearing short lat-
eral setule, spine next medial longer than setule,
midlateral setules diverse. Cuticle with ordinary
bulbar setules.
OBSERVATIONS.?Juvenile differing from adult
holotype mainly in smaller eyes and lack of dorsal
spine on outer ramus of uropod 2.
ILLUSTRATIONS.?Head heavily restored; upper lip
and maxilla 2 damaged during dissection, maxilli-
ped of holotype missing, these appendages taken
from juvenile; upper lip and epistome from ante-
rior view similar to illustration shown for B. myal-
lus (4) but epistomal region slightly smaller; max-
illa 2 like that of B. myallus but medial setae of
inner plate absent; maxilliped as shown for B.
myallus but inner plate with only 2 thick apical
spines, outer plate lacking lateral setae but palp
article 4 with 2 setules at base of nail; maxilla 1
as shown for B. panamunus (I), inner plate with 2
large and 2 small setae, apex of palp with 4 thick
and 4 thin setae (from holotype); peduncle of uro-
pod 3 shriveled and restored in drawing; setule on
coxa 7 missing; one spine on outer ramus of uro-
pod 3 missing.
HOLOTYPE.?NMV, female "a," 4.3 mm.
TYPE-LOCALITY.?PPBES 986/1, 12 Oct 1971,
Port Phillip Bay. Victoria, Australia, 4 m, sand.
VOUCHER MATERIAL.?Type-locality, juvenile "b,"
4.1 mm. Male unknown.
RELATIONSHIP.?This species, furnished with an
excavate posterodorsal margin of pleonite 3 and
lateral shield on the urosome, has the appearance
of a being a superadult of B. muldarpus (9), but
differs markedly in the much longer wrists on the
gnathopods, thinner hands and more transversely
oriented palms.
Birubius thalmus differs from B. cartoo (7) in
the smaller tooth of epimeron 3, simple cuticular
setules lacking alate plaques and the longer article
2 of antenna 1.
MATERIAL.?PPBES, one sample from one sta-
tion (2).
DISTRIBUTION.?Victoria, Port Phillip Bay, 4 m,
sand.
9. Birubius muldarpus, new species
FIGURES 110-112
DESCRIPTION OF FEMALE.?Head about 21 per-
cent of total body length, greatest width about 60
percent of length; rostrum slightly constricted, nar-
row, not reaching middle of article 2 on antenna 1.
Eyes large, largely occluded with pigment. Article
1 on peduncle of antenna 1 about 1.3 times as long
as wide, about twice as wide as article 2, ventral
margin with about 11 setules, weakly produced
dorsal apex with 3 setules; article 2 about 0.73
times as long as article 1, with ventral cycle of 7
setae; primary flagellum with 9-10 articles, about
as long as peduncle, lacking aesthetascs; accessory
NUMBER 345 243
S
FIGURE 110.?Birubius muldarpus, new species, holotype, female "a," 3.50 mm (n ? male "n,1
3.64 mm).
244 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
flagellum with 7-8 articles. Spine formula on arti-
cle 4 of antenna 2 = 1-3-4-3, dorsal margin with
notch bearing 3 setae, ventral margin with 4-5
groups of 1-4 long to medium setae, one ventro-
distal long spine; article 5 about 0.75 times as long
as article 4, facial spine formula = 1-2-1, 1-2-0, or
1-2-2, dorsal margin naked, ventral margin with
3 sets of 1-2 long to short setae, 2 ventrodistal long
to medium spines and setae; flagellum about 1.4
times as long as articles 4-5 of peduncle combined,
with 11 articles. Mandibles with medium palpar
hump; right incisor with 3 teeth and accessory
tooth; left incisor with 4 humps in 2 branches;
right lacinia mobilis bifid, distal branch little
shorter than proximal, distal branch flabellate, den-
ticulate, proximal branch simple, pointed, with 2
facial humps; left lacinia mobilis with 4 teeth,
middle teeth shortened; right rakers 9, plus one
rudimentary; left rakers 9; molar in form of bul-
bous hump demarcated mainly by spines, molars
with 4-5 primarily long spines, plus one weakly
disjunct, each molar with plume; palp article 1
short, article 2 with one short inner apical seta
and 2 other shorter inner setae, article 3 about as
long as article 2, oblique apex with 9 spine-setae,
basofacial formula = 1 + 1. Inner plate of maxilla
1 ordinary, bearing one long apical pluseta, one
similar apicomedial seta, 2 apicolateral much
shorter setae; palp article 2 with 4 apicomedial
marginal spines and 3 submarginal setae. Plates of
maxilla 2 extending equally, outer slightly broader
than inner, outer with 3 apicolateral setae, inner
with one medial seta. Inner plate of maxilliped
with 2 large, thick, apical spines, 3 apicofacial
setae, 3 medial setae; outer plate with 6 medial and
apical spines, 2 apicolateral setae; palp article 1
with one apicolateral seta, article 2 with 4 apico-
lateral setae, medial margin of article 2 moderately
setose, article 3 weakly protuberant, with 3 facial
setae, 2 lateral setae, nail of article 4 long to me-
dium, with 2 accessory setules. Coxa 1 expanded
distally, anterior margin weakly concave; main ven-
tral setae of coxae 1-4 = (5-7)-(5-7)- (4-7)-O, pos-
teriormost seta of coxae 1-2 shortest (or aberrantly
long on coxa 3); anterior and posterior margins of
coxa 4 almost parallel, posterior margin convex,
posterodorsal corner sharply rounded, posterodor-
sal margin short, concave, width-length ratio of
coxa 4 = 19:22; long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 ? (1?2)?3?
(2-3)-3, short anteriors = (2-3)-(2-3)-(2-3)-(l-2),
long anteriors = (2-4)-(5-6)-O-O, no others. Gnath-
opods ordinary; width ratios of articles 5-6 on
gnathopods 1-2 = 26:35 and 26:35, length ra-
tios = 67:63 and 60:58; palmar humps small,
palms oblique; article 5 of gnathopod 1 elongate,
ovate, posterior margin almost flat; article 5 of
gnathopod 2 ovate, posterior margin almost flat.
Pereopods 1-2 similar; facial setae formula on arti-
cle 4 = 3 and 3, on article 5 = 4 and 5; main spine
of article 5 extending to M. 80 on article 6, article
5 with 2 proximoposterior spines; spine formula of
article 6 = 3-4 and 1-5 plus iniddistal seta; accliv-
ity on inner margin of dactyls of perco|>ods 1-2
sharp, produced as tooth, emergent setule long,
midfacial pluseta short. Coxae 5-7 posteroventral
setule formula 1-1-1. Articles 4-5 of j>ercopods
3-4 medium broad, facial spine rows moderately
developed, facial ridge formula on article 2 of
pereopods 3-5 = 0-2-2; width ratios of articles 2,
4, 5, 6 of pereopod 3 = 41:40:35:17, of pereopod
4 = 64:37:25:13. of pereopod 5-78:24:22:11,
length ratios of pereopod 3 = 70:27:36:41, of
pereopod 4 = 80:47:37:50, of pereopod 5 = 100:
26:21:27; article 2 of pereopod 5 slightly exceed-
ing apex of article 4; medial apex of article 6 bear-
ing 5 digital processes. Posteroventral corner of
epimeron 1 rounded, posterior margin weakly con-
vex, anteroventral margin with 2-4 medium setae,
posteroventral face with 2 long setae; posteroven-
tral corner of epimeron 2 weakly protuberant, pos-
terior margin convex, facial setae = 7-9; postero-
ventral corner of epimeron 3 weakly protuberant,
with setule sinus, posterior margin almost straight,
face with crescent of 3-6 setae near ventral poste-
rior margin; epimeron 3 with setule on postero-
dorsal margin. Urosomite 1 naked, articulation line
complete; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1-2 with articulate enlarged api-
cal nails, inner ramus of uropod 1 with two acces-
sory nails, outer ramus of uropod 1 with 3-4 dorsal
spines, inner with one, outer ramus of uropod 2
with 2 dorsal spines, inner with one circular dorso-
medial spine; peduncle of uropod 1 with 2 apico-
lateral spines and 2 basofacial setae, medially with
3 marginal setae and spines, apicalmost an enlarged
spine; peduncle of uropod 2 with 5-7 dorsal spines,
medially with one small apical spine. Peduncle of
uropod 3 with 6 ventral spines, dorsally with one
lateral spine, one medial setule; rami submascu-
NUMBER 245 245
FIGURE 111.?Birubius muldarpus, new species, holotype, female "a," 3.50 mm (c = female "c,1
3.39 mm; n ? male "n," 3.64 mm).
246 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 112-?Birubius muldarpus, new species, holotype, female "a," 3-50 mm (n ? male "n,"
3.64 mm).
line, inner extending to M. 80 on article 1 of outer
ramus, apex with 2 setae, medial margin with 3
setae, lateral margin naked, article 2 of outer ramus
short, 0.14, bearing 2 long setae, medial margin of
article 1 with 2 setae, lateral margin with 3 acclivi-
ties, spine formula = 1-1-1-1 or 2-1-1-1, setal for-
mula = 1-1-1-2 or 0-1-1-1. Telson ordinary,
length-width ratio = 5:4, not fully cleft, each apex
wide, rounded, lateral acclivity shallow, weak, bear-
ing long lateral setule, spine next medial shorter
than setule, midlateral setules equal on holotype,
diverse in other specimens. Cuticle with ordinary
bulbar setules mixed with pipes, surface lacking
fine striations.
OBSERVATIONS (female).?Female "c": Setal for-
mulae of article 4 on pereopods 1-2 = 3 and 3, of
article 4 = 4 and 5, spine rows of article 6 = 3 and
5 plus seta; this specimen better developed than
holotype except in outer ramus of uropod 3 and
peduncle of uropod 2, the high counts of spines
and setae on various appendages in the description
mainly originating from this specimen.
DESCRIPTION OF MALE.?Rostrum slightly longer
and narrower than in female. Article 2 of antenna
NUMBER 245 247
1 with 5 ventral setae; primary flagellum with 10
articles, one calceolus each on articles 1-6, aes-
thetascs poorly developed or absent; accessory fla-
gellum with 7 articles. Facial spine formula on
article 4 of antenna 2 = 0-3-3-3, on article 5 = 2,
article 5 with 3 dorsal sets of male setae and 2 cal-
ceoli, ventrodistal apex with 2 thin spines; flagellar
formula = 26, 2, 3, 5, 7 . . . 25. Right rakers 8 plus
1-2 rudimentaries; left rakers 9; molars with 6
spines plus one disjunct spine more strongly sepa-
rated than in female; basofacial setal formula of
article 3 on mandibular palp = 2-1. Palp of max-
illa 1 with 5 spines and 4 setae. Palp article 3 of
maxilliped with 4 facial setae. Coxa 4 like that of
female but smaller in relation to coxa 1; ventral
setal formula of coxae 1-4 = (6-7)-6-5-0. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 1?(1?2)?2?2, long anteriors = 4-
(5-6)-0-0, short anteriors = 2-2-2-2, no others.
Facial and setal spine formulas of pereopods 1-2
on article 4 = 4 and 3, on article 5 = 5 and 5, on
article 6 = 3 + 5 plus middistal seta. Article 2 of
only pereopod 5 narrower than in female, pereo-
pod 4 slightly larger relative to pereopod 3 than
in female, pereopod 5 much larger, pereopod 5
with special spine on article 5. Epimera 1-3 broad-
ened, posterior margin of epimeron 3 shortened;
setal formulas, epimeron 1 anteroventral = 3, pos-
teroventral = 2, epimeron 2 facial = 6, epimeron
3 posterior = 2 setules, facial = 6. Spine formulas
of uropods, uropod 1 peduncle apicolateral = 3,
basofacial = 4, medially = 4 spines and setae, uro-
pod 2 peduncle dorsal = 13; dorsal spines on outer
ramus of uropod 1 = 4, of uropod 2 = 3, inner ramus
of uropod 1 = 1, of uropod 2 = 1 , ventral spines on
peduncle of uropod 3 = 7, spine formula on article
1 of outer ramus = 1-1-2-2-2-2, setal formula =
1-1-1-1-1-1. Telson not elongate, distal spines not
shortened.
ILLUSTRATIONS.?Palp of maxilla 1 fully flattened
in illustration; epimeron 1 of male heavily dam-
aged and illustration omitted.
HOLOTYPE.?AM, female "a," 3.50 mm.
TYPE-LOCALITY.?SBS D1S4, 17 May 1972, oft
Malabar, New South Wales, Australia, 31 m, coarse
sand.
VOUCHER MATERIAL.?Type-Locality: juvenile
"b," 2.70 mm. SBS D2S5: female "c," 3.39 mm
(illus.). EBS 20: male "n," 3.64 mm (illus.); female
"e," 2.97 mm.
RELATIONSHIP.?From the B. cartoo-thalmus
(7-8) group, B. muldarpus differs in the smaller
tooth and absence of midposterior setules on epi-
meron 3, the absence of a midfacial ridge on article
2 of pereopod 3 and the shorter wrists of the
gnathopods.
Birubius muldarpus differs from the B. myallus-
kareus (4-5) group in the absence of fully facial
setae on epimeron 3 and from B. myallus in addi-
tion by the smaller protrusion on epimeron 3.
From B. panamunus (1) and B. apart (6), B. mul-
darpus differs in the smaller tooth on epimeron 3
and the absence of a midfacial row of setae on
epimeron 3.
The absence of long setae on coxa 4 differen-
tiates B. muldarpus from B. lorus (2) and B. nam-
muldus (3).
Birubius yorlunus (31) has a simple right lacinia
mobilis and fully facial setae on epimeron 3.
See comments with B. thalmus about the possi-
bility of conspecificity.
MATERIAL.?SBS, 4 samples (4); EBS, one sample
DISTRIBUTION.?New South Wales: off Malabar,
31^19 m, sand and shell frags.; Jervis Bay, Mur-
ray's Beach, 6 m, Zostera.
10. Birubius gallangus, new species
FIGURES 113, 114
DESCRIPTION OF MALE.?Head about 15 percent
of total body length, greatest width about 70 per-
cent of length; rostrum constricted, short, apex
narrow, reaching almost to middle of article 2 on
antenna 1. Eyes medium, clear of opaque pigment,
but eosin colored. Article 1 on peduncle of an-
tenna 1 about 0.90 times as long as wide, about 2.5
times as wide as article 2, ventral margin with
about 12 setules, weakly produced dorsal apex with
2-3 setules, medial face with patch of fuzz; article
2 about as long as article 1, with ventral cycle of
4 setae; primary flagellum with 9 articles, about
0.85 times as long as peduncle, bearing one cal-
ceolus each on articles 1-4, no aesthetascs; acces-
sory flagellum with 6 articles. Spine formula on
article 4 of antenna 2 = 3-4?3, dorsomedial margin
fuzzy, ventral margin with 3 groups of 2 medium
setae, one ventrodistal medium spine; article 5
about 0.9 times as long as article 4, facial spine
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 113.?Birubius gallangus, new species, holotype, male "a," 3.26 mm (n ? male "n," 3.14
mm; q ? male "q," 3.50 mm; y = male "y," 3.11 mm; b ? epistome flat, z ? upper lip flat) .
NUMBER 245 249
formula = 2, dorsal margin bearing 3 sets of male
setae and 2 calceoli, ventral margin with 2 sets of
1-2 short to medium setae, 2 ventrodistal spines
and setule; flagellum elongate, flagellar formula =
(23-24), 1-3, 5, 7, . . . 23 or 2-4, 6, 8 . . . 20. Man-
dibles with strong palpar hump; right incisor with
3 teeth and accessory notch; left incisor with 4
humps in 2 branches; right lacinia mobilis bifid,
distal branch much shorter than proximal, weakly
expanded but pointed, proximal branch simple,
pointed, with marginal denticles; left lacinia mo-
bilis with 4 teeth; right rakers 5-7, plus rudimen-
taries; left rakers 6-7 plus 0-1 rudimentary; molar
in form of elongate plaque, right and left molars
with 5-6 primarily long spines plus one short spine
strongly disjunct; palp article 1 slightly elongate,
article 2 with one long inner apical seta and 2
other shorter inner setae, article 3 about 0.9 times
as long as article 2, oblique apex with 6 spine-
setae, basofacial formula = 0-1. Inner plate of max-
illa 1 ordinary, bearing one long apical pluseta,
one similar medial seta, 2-3 apicolateral much
shorter setae; palp article 2 with 4 apicalmedial
marginal spines and 3 submarginal setae. Plates of
maxilla 2 extending equally, outer slightly nar-
rower than inner, with 3 apicolateral setae, inner
with 2 medial setae. Inner plate of maxilliped with
one short apical spine set in middle (unusual), 5
mostly apicofacial setae; outer plate with 4 medial
and apical spines, no apicolateral setae; palp arti-
cles 1-2 each with apicolateral seta, medial margin
of article 2 moderately setose, article 3 protuberant,
with 2 facial setae, one lateral seta, nail of article
4 medium, with 2 accessory setules. Coxa 1 ex-
panded distally, anterior margin almost straight;
main ventral setae of coxae 1-4 = 6-6-3-0 and
0?(1?3)?(1?3)?1 rudimentaries, posteriormost seta
of coxae 1-2 shortened; anterior and posterior mar-
gins of coxa 4 divergent, posterior margin almost
straight, posterodorsal corner subsharp, posterodor-
sal margin long, weakly concave, width-length
ratio of coxa 4 = 17:19. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
1-2-3-3, others not noted. Gnathopods ordinary;
width ratios of articles 5-6 on gnathopods 1-2 =
28:36 and 28:39, length ratios = 72:75 and
66:65; palmar humps ordinary, palms oblique;
article 5 of gnathopod 1 elongate, ovate, posterior
margin rounded-flat; article 5 of gnathopod 2 elon-
gate, ovate, posterior margin rounded. Pereopods
1-2 similar, facial setae formula on article 4 = 4
and 4, on article 5 = 4 and 4; main spine of article
5 extending to M. 100 on article 6, article 5 with
one proximoposterior spine; spine formula of arti-
cle 6 = 3 + 3 plus middistal seta; acclivity on inner
margin of dactyls of pereopods 1-2 weak, obsoles-
cent, emergent setule long, midfacial pluseta highly
distad. Coxae 5-7 posteroventral setule formula =
1-1-1. Articles 4-5 of pereopods 3-4 of ordinary
width, facial spine rows sparse, facial ridge formula
on article 2 of pereopods 3-5 = 1-2-2, width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 47:38:34:16,
of pereopod 4 = 61:38:27:11, of pereopod 5 =
80:20:17:8, length ratios of pereopod 3 = 72:24:
35:40, of pereopod 4 = 8042:39:46, of pereopod
5 = 107=27:24:23; article 2 of pereopod 5 almost
reaching apex of article 4; medial apex of article 6
bearing digital processes. Posteroventral corner of
epimeron 1 rounded, posterior margin deeply con-
vex, anteroventral margin with 3 long setae, postero-
ventral face with 3 long setae; posteroventral corner
of epimeron 2 rounded-quadrate, posterior margin
strongly convex, facial setae = 6-7 crowded ante-
riorly; posteroventral corner of epimeron 3 with
small-medium upturned tooth with setule sinus,
posterior margin convex, with 1-2 setule notches,
posteroventral face with 1-2 setae; epimera 1-3
with small setule on posterodorsal margin set in
weak notch. Urosomite 1 with ventrolateral setule
at base of uropod 1, articulation line complete;
urosomite 3 weakly protuberant dorsally. Rami of
uropods 1-2 with articulate enlarged apical nails,
inner ram us of uropod 1 with one accessory nail,
outer ramus of uropod 1 with 2-3 dorsal spines,
inner with one, outer ramus of uropod 2 with 1-2
dorsal spines, inner with one dorsomedial spine;
peduncle of uropod 1 with 3 apicolateral spines,
basofacial setae absent, medially with 0-1 marginal
enlarged spine; peduncle of uropod 2 with 7-8
dorsal spines, medially with one medium apical
spine. Peduncle of uropod 3 with 4 ventral spines,
dorsally with one lateral spine, one medial spine;
rami masculine, inner extending to M. 110 on arti-
cle 1 of outer ramus, apex with 2 setae, medial and
lateral margins setose, article 2 of outer ramus
short, 0.13, bearing 2 long setae, medial margin of
article 1 setose, lateral margin with 4 acclivities,
spine formula = 1-1-2-2-2, setal formula = 1-1-1-
1-1. Telson elongate, length-width ratio = 7:5,
fully cleft, each apex protrusive, rounded, lateral
250
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 114.?Birubius gaUangus, new species, holotype, male "a," 3.26 mm.
NUMBER 245 251
acclivity deep, bearing ordinary lateral setule, spine
next medial slightly shorter than setule, midlateral
setules scarcely diverse, each lobe with dorsal row of
denticles. Cuticle with very sparse ordinary bulbar
setules.
VARIATIONS.?Head of male "q" with almost
blunt, poorly protuberant preocular corner (see
illustration); male "y" lacking distal branch on
right lacinia mobilis (see illustration), left lacinia
mobilis only trifid; left lacinia mobilis of male "n"
poorly 4-dentate; male "q" with spine formula on
article 4 of antenna 2 = 3-5-3 (otherwise males
with 3-4-3), 4 apicolateral spines on peduncle of
uropod 1, outer ramus of uropod 2 with 2 spines in
3 specimens, facial setae of epimeron 2 arranged in
zigzag fashion.
ILLUSTRATIONS.?Dorsal view of head of male "y"
bearing only outline of eyes; palp of maxilla 1 as
illustrated, not flattened, and normal basal cusp
proximal to palp not shown; setule on coxa 6 and
epimeron 2 missing.
HOLOTYPE.?AM, male "a," 3.26 mm.
TYPE-LOCALITY.?AM P. 18125, 12 Dec 1939,
Antechamber Bay, Kangaroo Island, South Austra-
lia, with light in net at night.
VOUCHER MATERIAL.?Type-locality: male "n,"
3.14 mm (illus.); male "q," 3.50 mm (illus); male
"y," 3.11 mm (illus.); male "e," 3.34 mm. Female
unknown.
RELATIONSHIP.?Birubius gallangus fits Key B of
Birubius in the simplicity of the distal branch on
the right lacinia mobilis, but the presence of an
accessory apical nail on uropod 1 suggests affinity
with species 1-9 in Key A in which the distal branch
of the right lacinia mobilis is complex and uropod 1
bears accessory nails. See remarks with B. lorus (2).
This species appears to have more characters in
common with B. muldarpus (9) than any other
species but differs from B. muldarpus in the re-
duced setation on epimeron 3, crowded setae on
epimeron 2, different positions and lengths of facial
ridges on pereopod 5, presence of a ridge on pereo-
pod 3, shorter rostrum and no true apical spine on
the inner plate of the maxilliped.
Birubius gallangus might have a point of origin
near B. panamunus (1) and B. cartoo (7) but differs
in the small tooth and poor setation of epimeron 3,
and the narrower rostrum. Birubius gallangus dif-
fers from the greater B. myallus-B. kareus (4-5)
group in the reduced setation and sharper tooth of
epimeron 3.
REMARKS.?The apical spine on the inner plate
of the maxillipeds is probably not homologous to
the spines normally occurring in other species and
therefore specific comparisons are made on the
basis that the normal apical spine is absent.
MATERIAL.?AM, one sample (40).
DISTRIBUTION.?South Australia, Kangaroo Is-
land, Antechamber Bay, neritic.
11. Birubius mayamayi, new species
FIGURES 115-118, U9(part)-121
DESCRIPTION OF FEMALE.?Head about 19 percent
of total body length, greatest width about 58-68
percent of length; rostrum unconstricted, broad,
reaching or exceeding middle of article 2 on an-
tenna 1. Eyes medium, largely occluded with pig-
ment. Article 1 on peduncle of antenna 1 about 1.3
times as long as wide, about twice as wide as article
2, ventral margin with about 10 setules, weakly pro-
duced dorsal apex with one setule; article 2 about
0.8 times as long as article 1, with ventral cycle of
6 setae; primary flagellum with 9-11 articles, about
0.90 times as long as peduncle, bearing several long
aesthetascs; accessory flagellum with 6-8 articles.
Spine formula on article 4 of antenna 2 = 1-3 4 4,
dorsal margin with notch bearing 3 setae, ventral
margin with 5-6 groups of 1-2 long and medium
setae, one ventrodistal long spine; article 5 about
0.8 times as long as article 4, facial spine formula =
1-2-2-2 or 1-2-3-2 or 1-2-2-1, dorsal margin bear-
ing 2 setules, ventral margin with 5 sets of 1-2
setae, 2 ventrodistal long spines, one subdistal facial
spine; flagellum about 1.3 times as long as articles
4-5 of peduncle combined, with 9-13 articles. Man-
dibles with weak to medium palpar hump; right
incisor with 3 teedi and notch; left incisor
with 4 teeth on 2 branches; right lacinia mobilis
bifid, distal branch much shorter than proximal,
distal branch flabellate, broad, variable, subbifid or
denticulate, occasionally with facial humps, proxi-
mal branch simple, pointed, with marginal denti-
cles; left lacinia mobilis with 4 teeth plus one acces-
sory tooth, middle teeth slightly shortened; right
rakers 8-10; left rakers 9-11; molar in form of
elongate plaque, right and left molars with 8-9
primarily long spines, each molar with plume; palp
article 1 short, article 2 with one long inner apical
252 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 115.?Birubius mayamayi, new species, holotype,
female "z," 4.78 mm (key to lower case letters in Figures 115?
117: b mm male "b," 4.60 mm; c ? male "c," 4.61 mm; d ?
male "d," 4.10 mm; e = male "e," 4.70 mm; / ? female "t"
4.2 mm; g ? female "g," 4.72 mm; h ? female "h," 4.75
mm; i ? female "i," 4.2 mm; / ? female "j," 4.40 mm; * ?
female "k," 4.50 mm; I ? female "1," 4.60 mm; m ? juvenile
"m," 3.50 mm; n ? juvenile "n," 3.50 mm) .
NUMBER 245 253
FIGURE 116.?Birubius mayamayi, new species (see legend for Figure 115).
254 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 117.?Birubius mayamayi, new species (see legend for Figure 115).
NUMBER 245 255
wS
FIGURE 118.?Birubius mayamayi, new species, female "a," 4.9 mm (w ? female "w," 4.3 mm;
y ? female "y," 4.65 mm; line near A2 ? length of remaining flagellum).
256
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Git
FIGURE 119.?Upper: Birubius mayamayi, new species, female "a," 4.9 mm (n/ ?? female "w," 4.S
mm). Lower: Birubius wirakus, new species, holotype, female "a," 4.3 mm.
NUMBER 245 257
X2
FIGURE 120.?Birubius mayamayi, new species, female "v," 4.92 mm (5 = female "s," 4.64 mm).
seta and 1-3 other similar or shorter inner setae,
article 3 about 1.3 times as long as article 2, oblique
apex with 8-9 spine-setae, basofacial formula = 1-2.
Inner plate of maxilla 1 ordinary, bearing one long
apical pluseta, one shorter similar apicomedial seta,
2 apicolateral much shorter setae; palp article 2
with 3-4 apicalmedial marginal spines and 3-5 sub-
marginal setae. Plates of maxilla 2 extending sub-
equally, of subequal breadth, outer with 3-4 apico-
lateral setae, inner with one medial seta. Inner
plate of maxilliped with 2 large thick apical spines,
2-3 apicofacial setae, 3-6 medial setae; outer plate
with 10-11 medial and apical spines, 4?6 apico-
lateral setae; palp article 1 with apicolateral seta,
article 2 with one apicolateral seta, medial margin
of article 2 moderately setose, article 3 not protu-
berant, with 4 facial setae, no lateral setae, nail of
article 4 long, with 2 accessory setules. Coxa 1 ex-
panded distally, anterior margin straight; main ven-
tral setae of coxae 1-4 = 9-9-7-11, posteriormost
258 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
seta of coxae 1-3 shortest; anterior and posterior
margins of coxa 4 strongly divergent, posterior
margin very oblique, weakly convex, posterodorsal
corner sharp, posterodorsal margin ordinary, weakly
V-shaped, width-length ratio of coxa 4 = 15:16.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = (2-3)-3-3-5, plus one long
facial on gnathopod 1, long anteriors = 4-(7-9)-O-O,
short anteriors = (1-2)- (l-3>-(l-2)-(l-2). Gnatho-
pods generally ordinary, hands slightly broadened;
width ratios of articles 5-6 on gnathopods 1-2 =
25:36 and 27:38, length ratios = 65:68 and 59:62;
palmar humps ordinary to small, palms oblique;
article 5 of gnathopod 1 elongate, ovate, posterior
margin rounded; article 5 of gnathopod 2 elongate,
ovate, posterior margin rounded. Pereopods 1-2
similar, facial setae formula on article 4 = 5 and 5,
on article 5 = 6 and 6 or 5 and 5; main spine of
article 5 extending to M. 95 on article 6, article 5
lacking proximoposterior spines; spine formula of
article 6 = 4 + 5 or 3 + 4 plus middistal seta, some
spines especially long; acclivity on inner margin of
dactyls of pereopods 1-2 weak, emergent setule
short, midfacial pluseta ordinary. Coxae 5-7 pos-
teroventral setule-seta formula = 7-3-1. Articles
4-5 of pereopods 3-4 of ordinary width, facial spine
rows sparse, facial ridge formula on article 2 of
pereopods 3-5 = 0-2-2; width ratios of articles 2,
4, 5, 6 of pereopod 3 = 40:35:29:15, of pereopod
4 = 57:25:17:8, of pereopod 5 = 80:22:20:7, length
ratios of pereopod 3 = 68:28:39:39, of pereopod
4 = 86:45:45:58, of pereopod 5 = 88:25:25:28;
article 2 of pereopod 5 almost reaching middle
of article 5; medial apex of article 6 finely
combed, bearing 5-6 digital processes. Posteroventral
corner of epimeron 1 weakly protuberant, posterior
margin weakly convex, with 0-3 setules, corner with
or without setule, anteroventral margin with 4-5
medium setae, posteroventral margin with 2 long
setae; posteroventral corner of epimeron 2 with
small rounded or pointed tooth guarded by setule
sinus, posterior margin weakly convex, with 2-3
setules, facial setae = 8-11, occasional pair set verti-
cally; posteroventral corner of epimeron 3 rounded,
weakly protuberant, with small rounded tooth
guarded by setule sinus, posterior margin straight,
with 3 setule notches, ventral margin with 2-4 setae
evenly spread; epimera 1-3 with large to small seta
on posterodorsal margin set in weak notch. Uroso-
mite 1 with ventral setule at base of uropod 1, artic-
ulation line complete; urosomite 3 unprotuberant
dorsally. Rami of uropods 1-2 with articulate en-
larged apical nails, rami of uropods 1-2 lacking
accessory nails, outer ramus of uropod 1 with 5-6
dorsal spines, inner with 2-3, outer ramus of uro-
pod 2 with 4 dorsal spines, inner with 1-2 dorso-
medial spines; peduncle of uropod 1 with 2 apico-
lateral spines and 4-5 basofacial setae, medially
with 5-6 marginal spines, apicalmost enlarged; pe-
duncle of uropod 2 with 7-9 dorsal spines, medially
with one apical spine. Peduncle of uropod 3 with
4-5 ventral spines, dorsally with one lateral spine,
2 medial setules or one spine and one setule; rami
submasculine, inner extending to M. 85 on article
1 of outer ramus, apex with 2 setae, medial and
lateral margins more or less setose, article 2 of outer
ramus short, 0.15-0.22. bearing 2 long setae, medial
margin of article 1 with 1-4 setae, lateral margin
with 4 acclivities, spine formula = 1-1-1-1-1, setal
formula = 1?1?1?1?2. Telson long, length-width
ratio = 12:11, almost fully cleft, each apex of me-
dium width, rounded, lateral acclivity broad, shal-
low, weak, bearing ordinary lateral setule, spine
next medial of length equal to or shorter than set-
ule, midlateral setules highly diverse. Cuticle with
sparse ordinary bulbar setules of varying sizes mixed
with pipes, emergent setules ordinary.
DESCRIPTION OF MALE.?Rostrum and head
slightly shorter and narrower than in female.
Primary flagellum of antenna 1 with 10-12 articles,
one calceolus each on articles 1-6; accessory flagel-
lum with 6-7 articles. Facial spine formula on arti-
cle 4 of antenna 2 = 3-4-4, of article 5 = 2-2-2 or
2-2, article 5 with 2 dorsal sets of male setae and
1-3 calceoli, ventrodistal apex with 3 setules, flagel-
lar formula = 28, 1-3, 5, 7 . . . 27. Article 2 of man-
dibular palp with 2-3 inner setae, basofacial setal
formula on article 3 = 2-2 or 3-2. Article 2 of only
pereopod 5 narrower than in female, article 6 and
dactyl of pereopod 3 elongate, articles 5-6 narrower
than in female, anterior facial ridge of pereopod 5
shortened, article 6 subequal to article 5 in length.
Epimera 1-3 broadened; posterior margin of epi-
meron 3 shortened, posterior setule notches more
crowded and sharper than in female; dorsoposterior
setule on epimera 1-2 absent, shortened on epi-
meron 3; setal formulas, epimeron 1 anteroventral
= 5, posteroventral = 2, epimeron 2 facial - 9-14,
epimeron 3 posterior = 4, ventral = 3-5. Spine
formulas of uropods, uropod 1 peduncle apico-
NUMBER 245 OHQ
FICURE 121.?Birubius mayamayi, new species, female "v," 4.92 mm.
260 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
lateral = 2-4, basofacial = 6-7, medial ?= 5, uropod
2 peduncle dorsal ? 12-16, dorsal spines on outer
ramus of uropod 1 = 6-7, of uropod 2 = 5-6, inner
ramus of uropod 1 = 2-3, of uropod 2 = 1-2.
Telson slightly elongate, distal spines shortened.
VARIATIONS.?The major variant, of subtle dis-
tinction, comprises specimens with shorter, apically
broader rostrum combined with slightly stouter arti-
cle 6 of pereopod 4, somewhat narrower hands of
gnathopods as illustrated and a spine instead of
setules on the apicomedial margin of uropod 3
peduncle. But minor variations occur among other
specimens in lengths of spines on article 6 of pereo-
pods 1-2, dorsal outline of head and wide variations
in setal and spine counts on epimera and uropods
and especially in the form of the right lacinia
mobilis (as illustrated). Loss of pigment occurs in
the eyes of various specimens, after prolonged ex-
posure to formaldehyde, especially in those of the
"major variant" noted above.
Botany Bay Variants: Five out of six specimens
from New South Wales in EBS 304 and 311, differ
from Victorian material in the absence of a dorso-
medial spine on the inner ramus of uropod 2; one
specimen, female "p" of EBS 311 bears this spine.
Female "v" of that sample is unusual in that the 2
posterofacial setae on epimeron 2 are set hori-
zontally, not vertically. Female "v" is illustrated as
voucher material to demonstrate that, despite minor
differences, the Botany Bay material belongs with
B. mayamayi. Unillustrated parts are described as
follows: setal and spine formula of articles 4, 5, 6 of
pereopod 2 = (4-5)-6-(4 + 4); head about 19 per-
cent of total body length; view of upper lip shown
from oblique dorsal aspect; coxae and mandibles as
in Victorian material; flagellum of antenna 2 with
12 articles, 1.45 times as long as articles 4-5 of
peduncle together. The posteroventral tooth on
epimeron 2 is smaller and sharper in Botany Bay
material than in the majority of Victorian speci-
mens.
Male "u" of EBS 304: Article 4 of antenna 2 with
spine formula of 3-4-3, and on article 5 = 2-2.
HOLOTYPE.?NMV, female "z," 4.78 mm.
TYPE-LOCALITY.?PPBES 902/5, 21 Sep 1970, Port
Phillip Bay, Victoria, Australia, 10 m, sandy silt.
VOUCHER MATERIAL.?Illustrated: PPBES 902/2:
male "b," 4.60 mm; female "g," 4.72 mm; female
"h," 4.75 mm. PPBES 977/4: male "c," 4.61 mm.
PPBES 977/2: female "f," 4.2 mm. PPBES 961/1:
male "d," 4.10 mm. PPBES 964/4: male "e," 4.70
mm. PPBES 983/5: female "i," 4.2 mm; female "j ,"
4.40 mm. PPBES 951/2: female "k," 4.50 mm.
PPBES 959/5: female "I," 160 mm. PPBES 938/5:
juvenile "m," 3.50 mm; juvenile "n," 3.30 mm.
CPBS 03N/2: female "a," 4.9 mm; female "w," 4.3
mm. CPBS C6/3: female "y." 4.65 mm. EBS 304:
female "v," 4.92 mm; female "s," 4.64 mm.
Not Illustrated: EBS 311: female #'p." 4.48 mm,
EBS 304: male "u." 3.94 mm.
RELATIONSHIP.?This species and species 12-15
differ from the B. panamunits group (1-10) in the
absence of accessory apical nails on uropod 1.
Birubius mayamayi forms the model to which
species 12-15 arc compared.
Because B. mayamayi retains an unconstricted
rostrum and bears a fully developed posterodorsal
setule on epimeron 3 the direct relationship to any
of species 1-10 appears to be remote. Only B.
panamunus (1) retains the unconstricted rostrum
but lacks the epimeral setule. Both B. mayamayi
and B. panamunus have setose coxa 4 but epimeron
3 of B. mayamayi is distinct in other ways, such as
reduction of tooth, loss of facial setae and develop-
ment of posteroventral setules.
MATERIAL.?PPBES, 104 samples from 43 stations
(169); CPBS, 5 samples from 5 stations (8); EBS, 11
samples from 11 stations (20).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 2-23 m, sand, clay, silty sand, sandy
silt, silty clay, clayey silt. New South Wales, Botany
Bay, Towra, 1-3 m, sand, Zoster a.
12. Birubius wirakus, new species
FIGURES 119 (part), 122
DESCRIPTION OF FEMALE.?Head about 21 percent
of total body length, greatest width about 66 per-
cent of length; rostrum weakly constricted, broad,
not reaching apex of article 2 on antenna 1. Eyes
small, weakly occluded with pigment or clear. Arti-
cle 1 on peduncle of antenna 1 about 1.6 times as
long as wide, about 1.7 times as wide as article
2, ventral margin with about 6 setules, weakly
produced dorsal apex with two setule-setae; arti-
cle 2 about 0.9 times as long as article 1, with
ventral cycle of 5 setae; primary flagellum with 9
articles, bearing several aesthetascs; accessory
flagellum with 7 articles. Spine formula on article 4
NUMBER 245 261
FIGURE 122.?Birubius wirakus, new species, holotype, female "a," 4.3 mm (n ? female "n,
S.76 mm).
262 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
of antenna 2 = 2-4-4 or 3-4-4, dorsal margin with
notch bearing 2 long and short setae, ventral
margin with 3 groups of 2-3 mainly long setae, one
ventrodistal long spine; article 5 about 0.75 times
as long as article 4, facial spine formula = 1-2,
dorsal margin naked, ventral margin with one set
of setae, ventrodistally with one medium spine, 2
long spines, one long seta, one setule, subdistal
facial spine; flagellum about 1.33 times as long as
articles 4-5 of peduncle combined, with 10 articles.
Mandibles with weak palpar hump; right incisor
with 3 teeth; left incisor with 4 humps in 2
branches; right lacinia mobilis bifid, distal branch
much shorter than proximal, distal branch broad,
subbifid, proximal branch simple, pointed, with
marginal denticles, humps; left lacinia mobilis with
4-5 teeth, middle teeth shortened; right rakers 6-8
plus one or more rudimentaries; left rakers 7-9
plus one or more rudimentaries; molar in form of
weak hump demarcated mainly by spines, right
molar with 5 primarily long spines plus one short
spine strongly disjunct, left molar with 4-5 primar-
ily long spines plus one short spine strongly dis-
junct (except on holotype), each molar with plume;
palp article 1 short, article 2 with one long inner
apical seta and 1-2 other short inner setae, article
3 about 1.1 times as long as article 2, oblique apex
with 6-8 spine-setae, basofacial formula = 0. Inner
plate of maxilla 1 large, bearing one long apical
pluseta, one shorter similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
3-4 apicalmedial spines and 3 submarginal setae.
Plates of maxilla 2 extending equally, outer slightly
broader than inner, outer with 3 apicolateral setae,
inner with one medial seta. Inner plate of maxil-
liped with 2 large thick apical spines, 3 apicofacial
setae, 3 medial setae; outer plate with 5-7 medial
and apical spines, one apicolateral seta; palp arti-
cles 1-2 lacking apicolateral setae, medial margin of
article 2 strongly setose, article 3 not protuberant,
with 3 facial setae, nail of article 4 long, with 2
accessory setules. Coxa 1 scarcely expanded distally,
anterior margin weakly convex, main ventral setae
of coxae 1-4 = 5-6-(5-6)-(4-5), posteriormost seta
of coxae 1-4 elongate; anterior and posterior
margins of coxa 4 almost parallel, posterior margin
oblique, almost straight, posterodorsal corner
rounded, posterodorsal margin weakly concave,
width-length ratio of coxa 4 = 17:20. Long poster-
ior setae on article 2 of gnathopods 1-2 and pereo-
pods i_2 = i_2-(0-2)-4, short anteriors =1-1-1-0,
long anteriors = 0-4-0-0, no others. Gnathopods
generally ordinary; width ratios of articles 5-6 on
gnathopods 1-2 = 25:34 and 2736, length ratios
= 70:65 and 61:64; palmar humps small, palms
weakly oblique; article 5 of gnathopod 1 weakly
elongate, ovate, posterior margin rounded-flat; arti-
cle 5 of gnathopod 2 ovate, posterior margin
rounded-flat. Pereopods 1-2 similar, facial setae
formula on article 4 = (2-4) and (2-3), on article
5 = 3 and 4; main spine of article 5 almost extend-
ing to M. 80 on article 6, article 5 with one proxi-
moposterior spine on pereopod 2 (rarely pereopod
1); spine formula of article f> = 4 + 4, rarely 4 + 5,
plus middistal seta, some spines especially long;
acclivity on inner margin of dactyls of pereopods
1-2 sharp, produced as tooth, emergent setule very
short, almost fully immersed, midfacial pluseta ordi-
nary. Coxae 5-7 posteroventral setule formula =
3-3-3. Articles 4-5 of pereopods 3-4 of ordinary
width, facial spine rows moderately developed,
facial ridge formula on article 2 of pereopods 5-5
= 0-2-2; width ratios of articles 2, 4, 5, 6 of pe-
reopod 3 = 39:37:34:17, of pereopod 4 = 58:32:
22:10, of pereopod 5 = 71:20:19:9, length ratios
of pereopod 3 = 64:25:34:34, of pereopod 4 =
71:43:38:50, of pereopod 5 = 80:22:22=25; article
2 of pereopod 5 reaching apex of article 4; me-
dial apex of article 6 weakly scalloped. Postero-
ventral corner of epimeron 1 narrowly rounded,
posterior margin weakly convex, with setule, antero-
ventral margin with 2-4 short setae, posteroventral
face with 1-2 long setae vertically paired; postero-
ventral corner of epimeron 2 subquadrate, with or
without setule, posterior margin almost straight,
facial setae = 5-6, occasionally posteriormost pair
set vertically; posteroventral corner of epimeron 3
weakly protuberant, with setule sinus, posterior mar-
gin almost straight, ventral margin with 2 setae;
epimera 1-3 with small to large seta on posterodorsal
margin set in weak to strong notch. Urosomite 1 with
lateral ventral setule at base of uropod 1, articula-
tion line almost complete; urosomite 3 unprotuber-
ant dorsally. Rami of uropods 1-2 with articulate
enlarged apical nails, rami of uropods 1-2 lacking
accessory nails, outer ramus of uropod 1 with 3
dorsal spines, inner with 2, outer ramus of uropod
2 with 2 dorsal spines, inner with one dorsomedial
spine; peduncle of uropod 1 with 2 apicolateral
spines and 2 basofacial setae, medially with 3
NUMBER 245 263
marginal spines, apicalmost enlarged; peduncle of
uropod 2 with 4 dorsal spines. Peduncle of uropod
3 with 4 ventral spines, dorsally with one lateral
spine, one medial spine; rami feminine, inner ex-
tending to M. 65 on article 1 of outer ramus, apex
with 1-2 short setae, article 2 of outer ramus
elongate, 0.31, bearing 2 setae, apicomedial margin
of article 1 with one long seta, lateral margin with
3 acclivities, spine formula = 1-1-1-1, setal formula
= 0-1-1-1. Telson ordinary, length-width ratio =
1:1, almost fully cleft, each apex wide, rounded,
lateral acclivity broad, weak, bearing ordinary lat-
eral setule, spine next medial much longer than
setule, midlateral setules diverse. Cuticle with ordi-
nary sparse bulbar setules.
OBSERVATIONS.?Female "n," 3.76 mm: Epimeron
2 more densely setose than in holotype (see illustra-
tion) but following features of weaker development:
spine formula of article 4 on antenna 2 (lesser count
stated in description), fewer setae on coxae 3-4, on
article 4 of pereopod 1, and on inner ramus of
uropod 3, setule count on coxae 5-7 = 4-2-2.
DESCRIPTION OF MALE (subadult, male "nn," 3.93
mm).?Scarcely distinct from female; eyes enlarged,
of medium size, darkly pigmented; facial spine
formula on article 4 of antenna 2 = 1-3-4-3, on
article 5 = 0-2, flagellum with 11 articles, basal
article stout and proliferating; disjunct spine on
right mandibular molar absent; formula of long
setae on coxae 1-4 = 6-6-6-3; length ratios of arti-
cles 5-6 on gnathopods 1-2 = 69:72 and 60:69; epi-
meron 1 with 5 anteroventral setae, posteroventral
face with 2 setae in tandem, epimeron 2 with 7
facial setae in tandem, posteriormost pair crowded
together; uropod 3 masculine, inner ramus slightly
exceeding apex of article 1 on outer ramus, apex
with 3 setae, outer ramus with 2 lateral acclivities,
setal formula = 1-1-2, spine formula = 1-1-1,
medial apex and margin with 2 setae in tandem,
peduncle with 5 ventral spines; main spine on each
lobe of telson shorter than in female; other char-
acters like female, especially right lacinia mobilis,
mandibular palp, antenna 1, dorsal margin of arti-
cle 4 on antenna 2, pereopods 1-5, epimeron 3,
uropods 1-2 except where mentioned, mouthparts
other than mandibles not examined.
DESCRIPTION OF JUVENILE (juvenile male "t," 3.32
mm).?Article 4 of antenna 2 with facial spine
formula of 1-3-3-1, article 5 with facial spine
formula of 0-2; both pereopods 1-2 with postero-
proximal spine on article 5 (unusual); epimeron 1
with 2 anteroventral setae, 2 posterofacial setae in
tandem, epimeron 2 with 8 facial setae, epimeron 3
with 2 ventral setae, posterior margin like adult but
with 3 setules in tandem just below dorsal seta in
crotch; outer ramus of uropod 1 with 2 dorsal
spines, of uropod 2 with one dorsal spine, peduncle
of uropod 2 with 3 dorsal spines; peduncle of uro-
pod 3 with 4 ventral spines, inner ramus of uropod
3 as long as article 1 of outer ramus, bearing 3
apical setae. Other external features like female;
mouthparts not examined.
ILLUSTRATIONS.?Species generally similar to
variant-phenotype of Birubius mayamayi (11) (fig-
ures 118, 119), and otherwise similar to main pheno-
type of that species, following illustrations therefore
omitted for B. wirakus: lateral head, antennae 1-2,
prebuccal mass, upper lip, palpar hump and palp of
mandibles, lower lip, maxillae 1-2, maxilliped,
urosome, coxae 1-3, cuticle. Following exceptions
for B. wirakus noted in description: spine formula
on article 5 of antenna 2, basofacial setae of man-
dibular palp article 3, spines and setae of maxil-
liped fewer, nail of dactyl of maxilliped more
elongate, uropodal spine formulas; view of left
mandible heavily flattened, with fourth molar spine
and smaller raker spines hidden in view as shown,
apex of that molar illustrated separately in 2 views.
Additional specific items not illustrated: midapex
on article 1 of antenna 1 with one setule, of article
2 with 3 setules, of article 3 with 4 apical and 2
ventral setules; article 3 of antenna 2 with one long,
one short facial setae, proximal facial spine groups
on article 4 of antenna 2 with accessory setule, with
2 subbasal penicillate setules on ventral margin,
article 5 of antenna 2 much narrower than article
4, dorsodistal apex with several setules; prebuccal
complex convex and moderately extended anter-
iorly, epistome and upper lip amalgamated, anter-
ior margin of upper lip with crescentic rugosity,
ventral margin truncate or weakly concave; lower
lip with one cone on outer lobe, mandibular lobes
rounded apically; outer plate of maxilla 1 with 11
spines, spine 2 broader; outer plate of maxilliped
with cusp at base of second apicomedial spine (but
absent on one side of holotype); inner ramus of
uropod 1 with normal inner basofacial seta; uro-
somites 1-2 amalgamated but articulation line com-
plete.
HOLOTYPE.?NMV, female, "a," 4.3 mm.
264 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TYPE-LOCALITY?CPBS B4/2, 14 Nov 1964, West-
ern Port, Victoria, Australia, 10.7 m, fine sand, mud.
VOUCHER MATERIAL.?PPBES 968/1, female "n,"
3.76 mm (illus.); EBS 32, male "nn," 3.93 mm;
EBS 31, juvenile male "t," 3.32 mm.
RELATIONSHIP.?This species closely resembles B.
mayamayi (11) and is difficult to distinguish except
on small characters requiring careful dissection and
observation. The main character of easily observ-
able condition is the presence of a stout proximo-
posterior spine on article 5 of pereopod 2, absent in
B. mayamayi. The following characters also dis-
tinguish B. wirakus from B. mayamayi: the blunter
cornered and squarer coxa 4, more robust pereopod
4, very slightly greater length of article 5 on
gnathopod 1 in female, absence of a posterodistal
tooth on epimeron 2 in female, longer article 2 on
outer ramus of uropod 3, presence of a disjunct
spine on molar in female, and proportional differ-
ences of articles on pereopods 3 and 5 (see illustra-
tions).
The holotype is a well advanced subadult female,
with brood lamellae almost fully grown but naked.
Immaturity is marked by poorly developed eyes,
paucity of setation, and feeble armament of uropod
3. A small number of subadults and juveniles of B.
mayamayi do have a shorter rostrum or a short arti-
cle 6 of pereopod 4 or both but detailed comparison
of B. wirakus with such subadults (of comparable
size and development) shows that specimens of B.
mayamayi have better developed eyes and more
heavily setose appendages. Their mandibles, coxae,
gnathopods and pereopods are virtually indis-
tinguishable from those of the terminal adult. They
lack a posteroproximal spine on article 5 of pereo-
pod 2, a characteristic of B. wirakus. The inner
ramus of uropod 3 is longer and article 2 of the
outer ramus is shorter in B. mayamayi.
MATERIAL.?PPBES, 2 samples (2); CPBS, one
sample (1); EBS, 2 samples (4).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 8-11 m, sand. New South Wales:
Jervis Bay, Murray's Beach, 6 m, Halophila.
13. Btrubhis chintoo, new species
FIGURES 123, 124
DESCRIPTION OF MALE.?Head about 20 percent
of total body length, greatest width about 70 per-
cent of length: rostrum weakly constricted, broad,
almost reaching middle of article 2 on antenna 1.
Eyes medium, clear of pigment, but eosin in color.
Article 1 on peduncle of antenna 1 about as long
as wide, about 2.6 times as wide as article 2, ventral
margin with numerous setules, medially with fuzz,
weakly produced dorsal apex with 2 setules; article
2 about as long as article 1, with ventral cycle of 5
setae (one hidden in illustration); primary flagellum
with 8 articles, about 0.9 times as long as peduncle,
bearing calceolus on articles 1-4; accessory flagellum
with 6 articles. Spine formula on article 4 of an-
tenna 2 = 1-2-4-3, dorsomedial margins of articles
3-4 fuzzy, ventral margin with 3-4 groups of 1-2
medium setae, one ventrodistal medium spine;
article 5 as long as article 4, facial spine for-
mula = 2, dorsal margin bearing 3 groups of male
setae, and 2 calceoli (one hidden in figure), posterior
margin with 2 sets of 3 setae and setules. 2 ventro-
distal short spines; flagellum elongate, flagellar
formula = (25-27), 2-7 . . . 21. 22 (calceolus on arti-
cle 23 rudimentary) or 2-4, 6, 8 . . . 26. Mandibles
with medium palpar hump; right incisor with 3
teeth and weak midhump; left incisor with 5 humps
in 2 branches; right lacinia mobilis bifid, distal
branch much shorter than proximal, weakly flabel-
late, subbifid of denticulate, proximal branch sim-
ple, pointed, with marginal denticles; left lacinia
mobilis with 4-5 teeth, middle teeth scarcely short-
ened; right and left rakers 8, plus tiny rudi-
mentaries; molar in form of elongate plaque, de-
marcated mainly by spines, right molar with 4-6
primarily long spines plus one short spine weakly
disjunct, left molar with 6 primarily long spines,
one of these scarcely disjunct, neither molar with
plume; palp article 1 short, article 2 with one long
inner apical seta and two other short inner setae,
article 3 about 1.1 times as long as article 2, oblique
apex with 7 spine-setae, basofacial formula = 0-2
or 0-1. Inner plate of maxilla 1 thin, bearing one
long apical pluseta, one similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
4 apicalmedial spines and 3 submarginal setae.
Plates of maxilla 2 extending equally, outer slightly
narrower than inner, with 2-3 apicolateral setae,
inner with one medial seta. Inner plate of maxil-
liped with 2 large, thick apical spines, 3 apicofacial
setae, 4 medial setae; outer plate with 7-8 medial
and apical spines, one apicolateral seta; palp article
2 with one apicolateral seta, medial margin of arti-
NUMBER 245 265
FIGURE 123.?Birubius chintoo, new species, holotype, male "a," 3.53 mm (6
S.61 mm).
male "b,
266 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
cle 2 weakly setose, article 3 scarcely protuberant,
with 2-3 facial setae, no lateral setae, nail of article
4 medium, with 2 accessory setules. Coxa 1 not ex-
panded distally, anterior margin straight; main
ventral setae of coxae 1-4 = 7-6-(5-6)-4, posterior-
most seta of coxae 1-3 shortened; anterior and
posterior margins of coxa 4 almost parallel, pos-
terior margin almost straight, posterodorsal corner
rounded, posterodorsal margin weakly concave,
width-length ratio of coxa 4 = 4:5. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 2-2-1-1, others not conspicuous or counted.
Gnathopods generally ordinary; width ratios of
articles 5-6 on gnathopods 1-2 = 26:34 and 27:35,
length ratios = 75:68 and 70:67; palmar humps
ordinary, palms oblique; article 5 of gnathopod 1
elongate, ovate, posterior margin flat; article 5 of
gnathopod 2 ovate, posterior margin flat. Pereopods
1-2 similar; facial setae formula on article 4 = 3-3,
on article 5 = 4-5; main spine of article 5 extend-
ing to M. 100 on article 6, article 5 with no proxi-
moposterior spines; spine formula of article 6 = 3-4
plus middistal seta, some spines especially long;
acclivity on inner margin of dactyls of pereopods
1-2 weak, sharp, emergent setule long, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule
formula = (2-3)-2-2. Articles 4-5 of pereopods 3-4
of ordinary width, facial spine rows sparse, facial
ridge formula on article 2 of pereopods 3?5 =
1-1-2; width ratios of articles 2, 4, 5, 6 of pereopod
3 = 44:36:37:18, of pereopod 4 = 66:37:23:11,
of pereopod 5 = 79:20:20:7, length ratios of pe-
reopod 3 = 77:30:40:40, of pereopod 4 = 88:56:
40:57, of pereopod 5 = 100:24:21:28; article 2 of
pereopod 5 reaching middle of article 4, article 5
with inner posterodistal spine of unusual shape,
granulate; medial apex of article 6 bearing weak
digital processes. Posteroventral corner of epimeron
1 rounded, posterior margin convex, corner with
setule notch, anteroventral margin with 6-7 medium
setae, posteroventral face with 2-3 long setae; postero-
ventral corner of epimeron 2 rounded, guarded by
setule sinus, posterior margin convex, facial setae
= 8-9; posteroventral corner of epimeron 3 weakly
protuberant, with setule sinus, posterior margin con-
vex, with one setule notch, ventral margin with 3
setae; epimera 1-2 with tiny seta on posterodorsal
margin set in weak notch. Urosomite 1 with ventral
setule at base of uropod 1, articulation line almost
complete; urosomite 3 unprotuberant dorsally.
Rami of uropods 1-2 with articlulate enlarged
apical nails, rami of uropods 1-2 lacking accessory
nails, outer ramus of uropod 1 with 3^i dorsal
spines, inner with 2, outer ramus of uropod 2 with
2-3 dorsal spines, inner with one dorsomedial spine;
peduncle of uropod 1 with 2 apicolateral spines and
2-3 basofacial setae, medially with 2 marginal setae
and apicalmost enlarged spine; peduncle of uropod
2 with 7-9 dorsal spines, medially with one tiny
apical seta. Peduncle of uropod 3 with 4 ventral
spines, dorsally with one lateral spine, one medial
spine; rami masculine, inner extending to M. 100
on article 1 of outer ramus, apex with 3 setae,
medial and lateral margins setose, article 2 of outer
ramus short. 0.18. bearing 2 long setae, apicomedial
margin of article 1 setose, lateral margin with 4
acclivities, spine formula 1-1-2-2-2. setal for-
mula = 1-1-1-1-1. Telson elongate, length-width
ratio =13:11, not fully cleft, each apex wide,
rounded, lateral acclivity weak, bearing ordinary
lateral setule, spine next medial shorter than setule,
each lobe with dorsal row of denticles, midlateral
setules diverse. Cuticle with ordinary sparse bulbar
setules.
ILLUSTRATIONS.?In dorsal view the head is out-
lined slightly broader than reality because of slight
squashing; ommatidia from dorsal view not differ-
entiated in the illustration; dorsal margin of eye
from lateral view slightly flattened owing to in-
ternal shrinkage and eye partially reconstructed by
reconsolidating one row of disjunct ommatidia.
HOLOTYPE.?AM, male "a," 3.53 mm.
TYPE-LOCALITY.?AM P. 18125, Antechamber Bay,
Kangaroo Island, South Australia, 12 Dec 1939,
with light in net at night.
VOUCHER MATERIAL.?Type-locality, male "b,"
3.61 mm (illus). Female unknown.
RELATIONSHIP.?This species differs from Biru-
bius mayamayi (11) in the absence of a dorsopos-
terior seta on epimeron 3, stronger bulge of epi-
meron 3; many fewer spines and setae on basofacial
uropod 1, outer rami of uropods 1-2 and peduncle
of uropod 2, all these spines larger; distinctive
shape of article 2 on pereopod 5, narrower articles
4-5 of pereopod 5; presence of only one pair of
spines on article 5 of antenna 2; larger pair of setae
on inner plate of maxilla 1 of equal size; presence
of a facial ridge on article 2 of pereopod 3, only
one ridge on pereopod 4; unexpanded coxa 1; and
stouter articles 4-5 of pereopods 3-4.
NUMBER 245 267
W
FIGURE 124.?Birubius chintoo, new species, holotype, male "a," S.53 mm.
268 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Only the slenderest of clues suggests that B.
chintoo is not the male of B. ivirakus (12): B. chin-
too has longer emergent setules on the dactyls of
pereopods 1-2, the facial ridge formulas on article
2 of pereopods 3-4 differ strikingly, the breadths
of the plates on maxilla 2 are different, the poster-
iormost seta on coxa 1-3 is shortened, and the
distal spine on article 5 of pereopods 1-2 is much
longer.
MATERIAL.?AM, one sample (9).
DISTRIBUTION.?South Australia, Kangaroo Island,
neritic.
14. Birubius karobrani, new species
FIGURES 125-127
DESCRIPTION OF FEMALE.?Head about 21 percent
of total body length, greatest width about 72 per-
cent of length; rostrum scarcely constricted, broad,
elongate, reaching middle of article 2 on antenna 1.
Eyes small, largely occluded with pigment. Article
1 on peduncle of antenna 1 about 1.4 times as long
as wide, about twice as wide as article 2, ventral
margin with about 10 setules, weakly produced
dorsal apex with 4 setules; article 2 about 0.84
times as long as article 1, with ventral row of 5
setae; primary flagellum with 11 articles, about 0.66
times as long as peduncle, bearing several short
aesthetascs; accessory flagellum with 8 articles.
Spine formula on article 4 of antenna 2 = 1-3-4-4,
dorsal margin with seta and notch bearing 2 setae,
ventral margin with 7 groups of 1-2 long and short
setae, one ventrodistal long spine; article 5 about
0.84 times as long as article 4, facial spine form-
ula = 1-2-2-2, dorsal margin naked, ventral mar-
gin with 4 sets of long to short setae, 2 ventrodistal
long to medium spines; flagellum broken, probably
exceeding length of articles 4-5 of peduncle com-
bined. Mandibles with weak palpar hump; right
incisor with 3 teeth and midnotch; left incisor with
5 teeth and humps on apical branch; right lacinia
mobilis bifid, distal branch much shorter than
proximal, flabellate, denticulate, proximal branch
simple, pointed, with marginal denticles; left
lacinia mobilis with 4 teeth plus one accessory
tooth, middle teeth shortened; right rakers 8 plus
one rudimentary; left rakers 9; molar in form of
elongate bulbous plaque, right molar with 9 pri-
marily long spines, one stub, left molar with 10
primarily long spines, each molar with plume;
palp article 1 short, article 2 with one short inner
apical seta and 2 other shorter inner setae, article S
about 1.3 times as long as article 2, oblique apex
with 9 spine-setae, basofacial formula = 1 - 1 . Inner
plate of maxilla 1 thin, bearing one long apical
pluseta, one shorter similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
5 apicalmedial marginal spines and 6 submarginal
or apical setae. Plates of maxilla 2 extending sub-
equally, outer scarcely broader than inner, outer
with 4 apicolateral setae, inner with no medial
setae. Inner plate of maxilliped with 2 large thick
apical spines, 2 apicofacial setae, 5 medial setae;
outer plate with 9 medial and apical spines, 4
apicolateral setae; palp article I with apicolateral
seta, article 2 with 2 apicolateral setae, medial mar-
gin of article 2 moderately setose, article 3 not pro-
tuberant, with 4 facial setae, no lateral setae, nail
of article 4 medium, with 2 accessory setules. Coxa
1 scarcely expanded distally, anterior margin con-
vex; main ventral setae of coxae 1-4 = 11-9-9-15,
posteriormost seta of coxae 1-4 shortest; anterior
and posterior margins of coxa 4 divergent, posterior
margin very oblique, convex, posterodorsal corner
subsharp, posterodorsal margin short, concave,
width-length ratio of coxa 4 = 9:8. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 2-3-2-3, long anteriors = (2-3)-9-O-O, short
anteriors = 3-3-4-2: no others. Gnathopods gen-
erally ordinary; width ratios of articles 5-6 on gna-
thopods 1-2 = 26:32 and 27:31, length ratios =
64:61 and 57:60, palmar humps ordinary, palms
oblique; article 5 of gnathopod 1 elongate, ovate,
posterior margin weakly rounded; article 5 of gna-
thopod 2 not elongate, ovate, posterior margin pro-
duced, flat. Pereopods 1-2 similar, facial setae
formula on article 4 = 6-6, on article 5 = 6-6;
main spine of article 5 extending to M. 100 on
article 6, article 5 lacking proximoposterior spines;
spine formula of article 6 = 4 + 5 and 5 + 5 plus
middistal seta, some spines especially long; acclivity
on inner margin of dactyls of pereopods 1-2 weak,
emergent setule short, midfacial pluseta ordinary.
Coxae 5-7 posteroventral setule formula = 5-2-1.
Articles 4-5 of pereopods 3-4 narrow, facial spine
rows sparse, facial ridge formula on article 2 of
pereopods 3-5 = 0-2-2, width ratios of articles 2, 4,
5, 6 of pereopod 3 = 39:28:26:11, of pereopod 4 =
56:28:21:11, of pereopod 5 = 77:21:17:7, length
NUMBER 245 269
FIGURE 125.?Birubius karobrani, new species, holotype, female "a," 5.88 mm.
270 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 126.?Birubius karobrani, new species, holotype, female "a," 5.88 mm.
ratios of pereopod 3 = 64:29:33:33, of pereopod
4 = 78:45:45:50, of pereopod 5 = 100:23:20:24; arti-
cle 2 of pereopod 5 reaching middle of article 5,
ventral margin deeply scalloped or toothed and bear-
ing long setae; medial apex of article 6 bearing 5
digital processes. Posteroventral corner of epimeron
1 quadrate, posterior margin straight, with 3 setae,
anteroventral margin with 4 short setae, postero-
ventral face with 3 setae, 2 anteriormost in vertical
pairing; posteroventral corner of epimeron 2 with
small sharp tooth, posterior margin weakly convex,
bearing 5 setae, facial setae = 1 1 , none in pairs set
vertically; posteroventral corner of epimeron 3 with
small tooth, posterior margin straight, 3 serrate,
with setae, ventral margin with 2 setae mainly
anteriad; epimeron 3 with 1-2 large setae on pos-
terodorsal margin. Urosomite 1 with articulation
line almost complete; urosomite 3 unprotuberant
NUMBER 245 271
FIGURE 127.?Birubius karobrani, new species, holotype, female "a," 5-88 mm (j ? juvenile
"j," 4.60 mm).
dorsally. Rami of uropods 1-2 with articulate en-
larged apical nails, outer ramus of uropod 1 with 4
dorsal spines, inner with 2, outer ramus of uropod
2 with 3 dorsal spines, inner with none; peduncle
of uropod 1 with 2 apicolateral spines and 8 baso-
facial setae, medially with 8 marginal setae and
spines, apicalmost an ordinary enlarged spine; pe-
duncle of uropod 2 with 6 dorsal spines, medially
with one medium apical spine. Peduncle of uropod
3 with 5 ventral spines, dorsally with one lateral
spine, one medial setule; rami feminine, inner ex-
tending to M. 80 on article 1 of outer ramus, apex
with one seta, medial and lateral margins naked,
article 2 of outer ramus short, 0.22, bearing 2 long
setae, apicomedial margin of article 1 with one
spine-seta, lateral margin with 3-4 acclivities, spine
formula = 1-1-1-2 or 1-1-1-1-2, setal formula =
1-1-1-0 or 0-1-1-1-1. Telson short, length-width
ratio =13:14, not fully cleft, each apex broad,
rounded, lateral acclivity broad, shallow, bearing
ordinary lateral setule, spine next medial longer
than setule, midlateral setules ordinary, diverse.
Cuticle with sparse ordinary bulbar setules, surface
bearing fine striations in form of linear fingerprint
pattern, emergent setules plumose.
OBSERVATIONS.?Juvenile "j," 4.60 mm: Differing
from described holotype female in much smaller
eye, presence of only 2 anteroventral setae on epi-
meron 1, 8 setae on epimeron 2, 3 posterior teeth
on epimeron 3, 3 dorsal spines on outer ramus of
uropod 1, 6 basofacial setae on peduncle of uropod
1, other external features fitting holotype, mouth-
parts not examined.
Crushed specimen WPBES 1732/1: Sex unknown,
female-like, differing from holotype in presence of
3 anteroventral setae on epimeron 1, 6 setae on epi-
meron 2, only one ventral seta on epimeron 3, outer
ramus of uropod 1 with 3 spines, 6 basofacial setae
on peduncle of uropod 1, 5 dorsal spines on pedun-
cle of uropod 2, other external features fitting holo-
type, mouthparts not examined.
ILLUSTRATIONS.?Pleonal view of epimeron 3 with
dotted extra posterodorsal seta present only on
right epimeron 3.
HOLOTYPE.?NMV, female "a," 5.88 mm.
TYPE-LOCALITY.?WPBES 1732/2, 26 Nov 1973,
Western Port, Victoria, Australia, 10 m, medium
sand.
VOUCHER MATERIAL.?Type-locality, juvenile "j,"
4.60 mm (illus.); WPBES 1732/1, 1 poor specimen.
Male unknown.
RELATIONSHIP.?This species is very close to but
apparently not the female of Birubius chintoo (13),
which is based only on the male. Birubius karobrani
272 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
differs from B. chintoo markedly in the long ventral
setae and teeth on article 2 of pereopod 5.
This species also differs from B. mayamayi (11)
and B. wirakus (12) in the teeth and setae of pereo-
pod 5, the elongation of the spine on the telson,
the presence of or larger size of the tooth on epi-
meron 2 and additionally, from B. wirakus, in the
less robust pereopod 4 and absence of stout spines
posteroproximally on article 5 of pereopods 1-2.
In the broadened coxa 4 and ventral setae on
article 2 of pereopod 5, B. karobrani appears to be
intermediate between the B. mayamayi group and
that centered by Parharpinia villosa and thus dem-
onstrates the convergence between these groups.
Birubius karobrani, however, differs from P. villosa
in numerous characters, among them: the normal
article 2 of pereopod 3 lacking posterior setae, the
normal formula of facial ridges on pereopods 3-5,
the unspiked cuticle, strong basofacial setal row on
uropod 1, absence of posterior setae on coxa 4, sim-
pler telsonic spination, longer article 3 of mandibu-
lar palp, distinct and articulate nail on the maxil-
lipedal dactyl, the absence of a disjunct molarial
spine, and the generic characters.
Birubius karobrani also has great interest be-
cause of its incipient characters of Heterophoxus,
such as weakly ensiform antenna 2 and ventrally
setose pereopod 5.
MATERIAL.?WPBES, 2 samples from one station
(3).
DISTRIBUTION.?Victoria, Western Port, 10 m,
sand.
15. Birubius booleus, new species
FIGURES 128-130
DESCRIPTION OF FEMALE.?Head about 19 percent
of total body length, greatest width about 65 per-
cent of length; rostrum very weakly constricted,
broad, reaching middle of article 2 on antenna 1.
Eyes medium, largely occluded with pigment. Arti-
cle 1 on peduncle of antenna 1 about 1.35 times
as long as wide, about twice as wide as article 2,
ventral margin with about 10 setules, weakly pro-
duced dorsal apex with 4 setules; article 2 about
0.85 times as long as article 1, with ventral cycle
of 7 setae; primary flagellum with 13 articles, about
0.9 times as long as peduncle, lacking aesthetascs;
accessory flagellum with 10 articles. Spine formula
on article 1 of antenna 2 = 1-3-4-4, dorsal margin
with notch bearing one seta and one setule, ventral
margin with 5 groups of 2-3 long to medium setae,
one ventrodistal medium spine; article 5 about 0.7
times as long as article 4, facial spine formula =
2-2, dorsal margin naked, ventral margin with 5
sets of 1-2 setae and setule, 3 ventrodistal long to
medium spines; flagellum about 1.3 times as long
as articles 4-5 of peduncle combined, with 13-14
articles. Mandibles with medium palpar hump;
right incisor with 3 teeth; left incisor with 2, distal
member bifid; right lacinia mobilis bifid, distal
branch little shorter than proximal, complex, ap-
parently composed of 2 fused raker spine-remnants
(see illustrations); left lat inia inobilis with 5 teeth,
middle teeth slightly shortened; right rakers 11 plus
one rudimentary; left rakers 13 plus one rudi-
mentary; molar composed of elongate plaque with
superimposed setulose bulbous hump, each with 7
long spines plus one shorter spine strongly disjunct,
each molar with plume; palp article 1 short, article
2 with 4 short inner setae and one other longer
apical seta, article 3 about as long as article 2,
oblique apex with 9 spine-setae, basofacial formula
= 0. Inner plate of maxilla 1 ordinary, bearing
one long apical pluseta, one similar apicomedial
seta, 2 apicolateral much shorter setae; palp article
2 with 4 apicomedial marginal spines and 4 sub-
marginal setae. Plates of maxilla 2 extending sub-
equally, outer broader than inner, with 4 apicolat-
eral setae; inner with 2 medial setae. Inner plate of
maxilliped with 2 large thick apical spines, 2 apico-
facial setae, 7 medial setae; outer plate with 7
medial and apical spines, 2 apicolateral setae; palp
article 2 with 2 apicolateral setae, medial margin of
article 2 moderately setose, article 3 protuberant,
with 4 facial setae, one lateral seta, nail of article 4
medium, with one accessory setule. Coxa 1 not ex-
panded distally, anterior margin straight; main
ventral setae of coxae 1-4 = 6-8-7-5, posteriormost
seta of coxae 1-3 shortest, one or two posterior
setae especially stout; anterior and posterior
margins of coxa 4 almost parallel, posterior margin
almost straight, posterodorsal corner rounded,
posterodorsal margin short, almost straight, width-
length ratio of coxa 4 = 17:21; long posterior setae
on article 2 of gnathopods 1-2 and pereopods 1-2
= 2-5-5-5, short anteriors = (4-6)-4-7-5, long
anteriors = 2-5-0-0, no others. Gnathopods gen-
erally ordinary; width ratios of articles 5-6 on
NUMBER 245 273
S
FIGURE 128.?Birubius booleus, new species, holotype, female "a," 6.37 mm.
274 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Dl
FIGURE 129.?Birubius booleus, new species, holotype, female "a," 6.37 mm.
gnathopods 1-2 = 23:35 and 25:37, length ratios =
78:60 and 67:60; palmar humps ordinary, palms
weakly oblique; article 5 of gnathopod 1 elongate,
posterior margin flat, long; article 5 of gnathopod
2 weakly elongate, posterior margin rounded-flat.
Pereopod 2 stouter than pereopod 1, especially arti-
cle 4, posterior margin of latter more strongly
setose, facial setae formula on article 4 = 4-3, on
article 5 = 5-6; main spine of article 5 extending
to M. 75 on article 6, article 5 with one proxi-
moposterior spine only on pereopod 2; spine
formula of article 6 = 5 + 6 plus tiny middistal
seta, some spines especially long; acclivity on inner
margin of dactyls of pereopods 1-2 sharp, produced
NUMBER 245 275
as tooth, emergent setule long, midfacial pluseta
ordinary. Coxae 5-7 posteroventral setule formula
= 3-4-4. Artciles 4-5 of pereopods 3-4 of ordinary
width, facial spine rows moderately developed,
facial ridge formula on article 2 of pereopods 3-5
= 0-2-2; width ratios of articles 2,4,5,6 of pereopod
3 = 47:40:35:15, of pereopod 4 = 72:30:22:11, of
pereopod 5 = 85:20:16:8, length ratios of pereo-
pod 3 = 71:36:40:40, of pereopod 4 = 83:55:49:52,
of pereopod 5 = 103:27=22:22; article 2 of pereopod
5 reaching middle of article 4; medial apex of arti-
cle 6 bearing 3 short digital processes. Posteroven-
tral corner of epimeron 1 rounded, posterior margin
weakly convex, corner with setule, anteroventral
margin with 11 short setae, posteroventral face with
2 long setae in vertical tandem; posteroventral
corner of epimeron 2 rounded or weakly protuber-
ant, guarded by weak setule sinus, posterior margin
almost straight, facial setae = 9, posteriormost pair
set vertically; posteroventral corner of epimeron 3
weakly protuberant, with setule sinus, posterior
margin straight, with 6 setule notches, ventral
margin with 2 setae; epimera 1-2 with setule on
posterodorsal margin set in weak notch. Urosomite
1 ventrally naked, articulation line almost com-
plete; urosomite 3 unprotuberant. Rami of uropods
1-2 with articulate enlarged apical nails, outer
ramus of uropod 1 with 5 dorsal spines, inner with
2, outer ramus of uropod 2 with 3-4 dorsal spines,
inner with 2 dorsomedial spines; peduncle of
uropod 1 with 3 apicolateral spines and 2 basofacial
small setules, medially with 6 marginal setae and
spines, apicalmost an ordinary spine; peduncle of
uropod 2 with 7 dorsal spines, medially with one
apical setule. Peduncle of uropod 3 with 6 ventral
spines, dorsally with one lateral spine, 2 medial
spines; rami submasculine, inner extending to M.
95 on article 1 of outer ramus, apex with one seta,
medial margin with 2, lateral margin with 4 setae,
article 2 of outer ramus short, 0.16, bearing 2 long
setae, medial margin of article 1 setose, lateral
margin with 4 acclivities, spine formula =
1?I?1?1?1, setal formula = 1-1-1-1-1. Telson ordi-
nary, length-width ratio = 1:1, almost fully deft,
each apex wide, rounded, lateral acclivity broad,
bearing ordinary lateral setule, spine next medial
much longer than setule, midlateral setules diverse,
small. Cuticle with ordinary bulbar setules of vary-
ing sizes closely packed with numerous large pipes,
emergent setules especially short.
OBSERVATIONS.?Lower lip like that of Birubius
chintoo (13) but extensively setulose; first raker on
left mandible peculiar (see illustration); apicalmost
facial spine on article 5 of antenna 2 so close to
distal spines as to be counted there.
HOLOTYPE.?NMV, female "a," 6.37 mm. Unique.
TYPE-LOCALITY.?WPBES 1735/2, 29 Nov 1973,
Western Port, Victoria, Australia, 9 m, muddy sand,
shell grit.
RELATIONSHIP.?The peculiar right lacinia
mobilis, apparently composed of two raker spines
fused together might place this species in the group
of Birubius (Key B) bearing a double-pronged right
lacinia mobilis but B. booleus bears only weak
resemblance to the other species in that group.
A closer resemblance occurs between B. booleus
and 3 species in the Birubius group with normal
lacinia mobilis but lacking accessory apical nails on
uropods 1-2: B. mayamayi (11), B. wirakus (12) and
B. chintoo (13). Birubius chintoo is based only on
males but is not the male of Cunmurra itickerus
because the latter has enlarged gnathopod 2.
Birubius booleus differs from the other 3 species
in the absence of basofacial setae on uropod 1 and
the distinctive right lacinia mobilis. In addition it
differs from B. mayamayi in the absence of a tooth
on epimeron 2, the greater spread of setae on
epimeron 2, the absence of posterior setae on epi-
meron 2, and in the vertical pairing of setae on
epimeron 1. It differs from B. wirakus, additionally,
in the absence of an enlarged posterodorsal seta on
epimeron 3. It further differs from B. chintoo in
the vertical pairing of setae on epimera 1-2, in the
disjunct spine on the molars and in the nonenlarged
spines on the inner ramus of uropod 1.
The very narrow anterior coxae suggest strong
affinity to Cunmurra so that B. booleus may ulti-
mately be separable generically from Birubius.
Birubius booleus could be distinguished from
Birubius on coxae 1-3, the right lacinia mobilis and
the somewhat reduced article 5 of antenna 2. How-
ever, the relationship to B. chintoo in epimeron 3
and coxa 1 is very strong and intergradations
towards other species of Birubius such as B. maya-
mayi suggest prudence in evaluation of the charac-
ters until further explorations of Australia reveal
additional evidence.
MATERIAL.?WPBES, one sample (1).
DISTRIBUTION.?Victoria, Western Port, 9 m,
muddy sand, shell grit.
276 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 130.?Birubius booleus, new species, holotype, female "a." 6-37 mm.
16. Birubius babaneekus, new species
FIGURES 131-133
DESCRIPTION OF FEMALE.?Head about 17 per-
cent of total body length, greatest width about 63
percent of length; rostrum weakly constricted, nar-
row, almost reaching middle of article 2 on antenna
1. Eyes small to medium, stained eosin color but
clear of occluding pigment. Article 1 on peduncle
of antenna 1 about 1.4 times as long as wide, about
twice as wide as article 2, ventral margin with
about 5 setules, weakly produced dorsal apex with
5 setules; article 2 about 0.7 times as long as article
1, with ventral cycle of 6-8 setae; primary flagellum
with 8-10 articles, about 0.75 times as long as
peduncle, bearing long aesthetascs; accessory
flagellum with 7 articles. Spine formula on article
4 of antenna 2 = 1-3-4-3 or 1-3-4-3-2, dorsal
margin with notch bearing 2-3 setae, ventral
margin with 4?5 groups of 1-2 long to medium
setae, one ventrodistal long spine; article 5 about
0.88 times as long as article 4, facial spine formula
= 1-2-2-2 or 1-2-2-2-1, dorsal margin with or
without setule, ventral margin with 4-5 sets of 1-2
long to short setae, 2 ventrodistal long spines;
flagellum about as long as articles 4-5 of peduncle
combined, with 8-10 articles. Mandibles with med-
ium palpar hump; right incisor with 3 teeth and
notch; left incisor with 4 humps in 2 branches;
right lacinia mobilis bifid, almost simple, distal
branch forming cusp on proximal branch, proximal
branch simple, pointed; left lacinia mobilis with 4
teeth, middle teeth shortened; right rakers 4?5; left
rakers 5-6 plus occasional rudimentary; molar in
form of elongate bulbous hump, each molar with 6
primarily medium spines, none disjunct, each with
plume; palp article 1 slightly elongate, article 2
with 1-2 long and medium inner apical setae and
2-3 other short inner setae, article 3 about 1.1 times
as long as article 2, oblique apex with 8 spine-setae,
basofacial formula = 1-2. Inner plate of maxilla 1
ordinary, bearing 4 short, similar setae; palp article
2 with 3-4 apicalmedial marginal spines and 3
submarginal setae. Plates of maxilla 2 extending
subequally, outer scarcely broader than inner, outer
with 3 apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with 2 large thick
apical spines, 2 apicofacial setae, 4-5 medial setae;
outer plate with 9 medial and apical spines, 4 api-
colateral setae; palp articles 1-2 with apicolateral
seta, medial margin of article 2 weakly setose, arti-
cle 3 unprotuberant, with 2-3 facial setae, one lat-
eral seta, nail of article 4 long, with 2 accessory
setules. Coxa 1 scarcely expanded distally, anterior
NUMBER 245 277
FIGURE 131.?Birubius babaneekus, new species, female "w," 3.9 mm (b ? male "b," 3.8 mm).
278 SMITHSONIAN CONTRIBUTION'S TO /OOLOGY
margin straight; main ventral setae of coxae 1-4 =
(9-lI)-(10-l 1)?(1O-13)-(13?17), posteriormost seta
of coxae 1-4 shortened; anterior and posterior
margins of coxa 4 slightly divergent, posterior
margin convex, posterodorsal corner sharp, slightly
protrusive, posterodorsal margin ordinary, flat,
width-length ratio of coxa 4 = 1:1. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1_2 = (3^)_(4_5)_(5_6)_(7_8), long anteriors =
(3-5)-(5-7)-O-O, others not counted. Gnathopods
ordinary, width ratios of articles 5-6 on gnathopods
1-2 = 31:37 and 30:37, length ratios = 63:71 and
56:65, palmar humps ordinary, palms oblique;
article 5 of gnathopod 1 ovate, posterior margin
rounded-flat; article 5 of gnathopod 2 shorter,
ovate, posterior margin rounded, short, almost
lobate. Pereopods 1-2 similar; facial setae formula
on article 4 = 6-8 and 7-8, on article 5 = 8 and 8;
main spine of article 5 extending to M. 100+ on
article 6, article 5 lacking proximoposterior spines;
spine formula of article 6 = 3 + 4 plus middistal
seta; acclivity on inner margin of dactyls of pereo-
pods 1-2 weak, emergent setule long, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule
formula = (3-4)-(3-4)-(l-2). Articles 4-5 of pereo-
pods 3?4 of ordinary width, facial spine rows sparse,
facial ridge formula on article 2 of pereopods 3-5
= 0-2-2; width ratios of articles 2, 4, 5, 6, of pereo-
pod 3 = 42:30:27:16, of pereopod 4 = 54:27:
19:9, of pereopod 5 = 76:19:16:7, length ratios
of pereopod 3 = 62:27:33:36, of pereopod 4 =
82:46:44:50, of pereopod 5 = 103:22:21:21; arti-
cle 2 of pereopod 5 reaching middle of article 5,
ventral margin with long setae; medial apex of arti-
cle 6 finely combed, bearing 5 digital processes.
Posteroventral corner of epimeron 1 rounded or
weakly protuberant, posterior margin convex, setose,
anteroventral margin with 3-5 medium setae, poster-
ventral face with one medium seta; posteroventral
corner of epimeron 2 protuberant, with small sharp
tooth, posterior margin convex, setose, facial setae =
7-11, occasional disjunct grouping anteriorly; pos-
teroventral corner of epimeron 3 with large upturned
tooth, posterior margin weakly convex, with 2 setae
near crotch, ventral margin with 6-11 setae mainly
in middle of margin. Urosomite 1 naked, articula-
tion line complete; urosomite 3 unprotuberant dor-
sally. Rami of uropods 1-2 with articulate enlarged
apical nails, outer ramus of uropod 1 with 5-8
dorsal spines, inner with 2, outer ramus of uropod
2 with 3 dorsal spines, inner lacking dorsomedial
spines; peduncle of uropod 1 with 1-2 apicolateral
spines and 6-8 basofacial setae, medially with 4-6
marginal setae and spines, plus apical enlarged
spine; peduncle of uropod 2 with 6-8 dorsal spines,
medially with one apical spine. Peduncle of uropod
3 with 4 ventral spines, dorsally with one lateral
spine, 2-3 medial setules; rami feminine, inner ex-
tending to M. 70 on article 1 of outer ramus, apex
with one seta, medial margin with one seta, article
2 of outer ramus short, 0.18. bearing 2 long setae,
apicomedial margin of article 1 with one seta, lat-
eral margin with 4 acclivities, spines long, for-
mula - 1-1-2-2-1 or 1-2-2-2-2. setal formula =
0-0-0-0-1. Telson ordinary, length-width ratio =
I:I, almost fully deft, each apex wide, rounded,
lateral acclivity deep, narrow, Ixarin^ ordinary lat-
eral setule, spine next medial shorter than or of
length equal to setule, occasionally with subapical
spine in middle of each lol>e, midlatcral setules di-
verse. Cuticle with ordinary bulbar setules, emerg-
ent setules short.
DESCRIPTION OF MALE.?Rostrum narrower than
in female. Primary flagellum of antenna 1 with
8-10 articles, one calceolus each on articles 1-5,
aesthetascs strongly developed; accessory flagellum
with 6 articles. Facial spine formula on article 4 of
antenna 2 = 1-3-4-3, on article 5 = 2-2, article 5
with 3 dorsal sets of male setae and one calceolus,
ventrodistal apex with 2 thin spines and one setule,
flagellar formula = (26-31), 1-6, 8, 10, . . . penulti-
mate, or 1-^ *, 6, 8 . . . or 2-5, 7, 9, . . . or 1, 3, 4, 6, 8
. . .. Coxa 4 narrow but smaller in relation to coxa 1
than in female; ventral setal formula of coxae
1-4 = fewer than in female. Facial and setal spine
formulas of pereopods 1-2 on article 4 = 6 and 6,
on article 5 = 6 and 6; articles 2 and 6 of pereopod
4 narrower than in female, posterior margin on
article 2 of pereopod 5 flattened. Epimeron 2 broad-
ened; posterior margin of epimeron 3 shortened;
setal formulas, epimeron 1 anteroventral = 4, pos-
teroventral = 1, epimeron 2 facial = 8 crowded,
epimeron 3 posterior = 1-2, tooth shortened, ven-
tral = 7. Spine formulas of uropods, uropod 1
basofacial = 6, medial = 4, uropod 2 peduncle
dorsal = 12-14, dorsal spines on outer ramus of
uropod 1 = 6, of uropod 2 = 4, inner ramus of uro-
pod 1 = 2, of uropod 2 = 1 , ventral spines on pe-
duncle of uropod 3 = 3, spine formula on article 1
of outer ramus = 0-0-1-1-1, setal formula = 2-1-
NUMBER 245 279
FIGURE 132.?Birubius babaneekus, new species, female "w," 3.9 mm (b ? male "b," 3.8 mm;
c " female "c," 4.8 mm).
280
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 133.?Birubius babaneekus, new species, female "w," 3.9 mm (a ?= holotype, female "a,"
3.9 mm; b = male "b," 3.8 mm; u = female "u," 4.0 mm; Y ?? venter of urosomite 1) .
1-1-1. Telson not elongate, distal spines not short-
ened.
OBSERVATIONS.?Spine formula on article 5 of
female antenna 2 commonly 1?3^?3 but next most
frequently occurring formulas = 1-3-4?3-1 or
1-3^-3-2, rarely 1-3-5-3 or 1-3-4-2, and one case
of 1-3-4-4-3; common spine formula on article 5 =
2-2-2, most frequent variant = 2-2-2-1 but one
abnormally large female, 5.0 mm, from CPBS 3IS/
468, has a formula of 2-2-2-2 on left antenna 2
and a normal formula of 2-2-2 on the right an-
tenna 2.
Ovigerous female "w," 3.9 mm, has an abnorm-
ally shaped coxa 7, partially fused inner lobes on
the lower lip, bears one sheathed terminal seta on
mandibular palp article 3, and an extra spine on
each telsonic lobe; the normal coxa 7 is illustrated
for the holotype.
One female, 4.5 mm, CPBS 32S/468, has greatly
shortened articles 5-6 on pereopod 4.
ILLUSTRATIONS.?Left mandible heavily flattened;
pereopodal dactyls as illustrated for B. cartoo (7)-
HOLOTYPE.?NMV, female "a," 3.9 mm.
TYPE-LOCALITY.?CPBS 32S/4, 12 Mar 1965,
NUMBER 245 281
Western Port, Victoria, Australia, 10 m, muddy
sand and gravel.
VOUCHER MATERIAL (all illustrated).?CPBS 3IS/
866, male "b," 3.8 mm; CPBS 200/5, female "c," 4.8
mm; CPBS 35S/5, female "w," 3.9 mm; PPBES
923/1, female "u," 4.0 mm.
RELATIONSHIP.?Three species, B. babaneekus, B.
gelarus (17) and B. quearus (18), form a group char-
acterized by: medium to large tooth on epimeron 3,
a right lacinia mobilis consisting of a simple spine
bearing a distomedial cusp, the absence of accessory
nails on uropods 1-2, presence of setae on coxa 4,
long to short but conspicuous ventral setae or set-
ules on article 2 of pereopod 5, broad, scarcely con-
stricted rostrum and strong basofacial setae on
uropod 1.
The large tooth of epimeron 3 and the setose
coxa 4 suggest affinities with B. panamunus (1) but
the absence of accessory nails on uropods 1-2 dif-
ferentiates the B. babaneekus group.
Certain affinity is shown to B. karobrani (14) in
the ventrally setose pereopod 5.
Species 20-27 to follow have a better developed
distal branch on the right lacinia mobilis but none
has the tooth of epimeron 3 as large as or as sharp.
The B. babaneekus group therefore appears to be
an independent cluster with unclear ancestry. The
unusual mandibular raker row has not been found
in any other group of phoxocephalids.
Birubius eleebanus (29) might be a descendant
of the B. babaneekus group but in addition to a
somewhat more normal raker row, B. eleebanus has
fewer posterior setae and a smaller tooth on epi-
meron 3.
MATERIAL.?CPBS, 41 samples from 15 stations
(60); WPBES, 2 samples from 2 stations (3); PPBES,
133 samples from 47 stations (297).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 2-25 m, silty clay, sand, clay, silty sand,
clayey silt.
17. Birubius gelarus, new species
FIGURES 134, 135
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about 70
percent of length; rostrum scarcely constricted,
broad, reaching middle of article 2 on antenna 1.
Eyes medium, stained eosin in color but clear of
occluding pigment, ommatidia especially small.
Article 1 on peduncle of antenna 1 about 1.3 times
as long as wide, about 2.3 times as wide as article 2,
ventral margin with about 9 setules, weakly pro-
duced dorsal apex with 4 setules; article 2 about
0.67 times as long as article 1, with ventral row of
3 setae; primary flagellum with 9 articles, about
0.73 times as long as peduncle, lacking aesthetascs;
accessory flagellum with 7 articles. Spine formula
on article 4 of antenna 2 = 1-3-4?3, dorsal margin
with notch bearing 2 setae and one spinule, ventral
margin with 6 groups of 2 long to short setae, one
ventrodistal long spine; article 5 about 0.85 times
as long as article 4, facial spine formula = 1-2-2-1
or 1-2-2-2 or 1-2-2, dorsal margin naked, ventral
margin with 4 sets of 1-2 long to short setae, 2
ventrodistal medium spines; flagellum about as
long as articles 4-5 of peduncle combined, with
9-10 articles. Mandibles with weak palpar hump;
right incisor with 3 teeth and notch; left incisor
with 6 humps in 2 weakly defined branches;
right lacinia mobilis weakly bifid, distal branch
much shorter than proximal, vestigial, proximal
branch simple, pointed, naked; left lacinia mobilis
with 4 teeth, middle teeth shortened; right rakers
4?5; left rakers 5-6; molar in form of elongate
plaque, right molar with 5-6 primarily medium
spines, one spine short and weakly disjunct, left
molar with 4-5 primarily long spines, one short
spine strongly disjunct, each molar with plume;
palp article 1 slightly elongate, article 2 with 1-2
long to medium inner apical setae and 2?4, rarely
one, other shorter inner setae, article 3 about 1.1
times as long as article 2, oblique apex with 8
spine-setae, basofacial formula = 1-2 or 0-1. Inner
plate of maxilla 1 ordinary to thin, bearing one
medium apical pluseta, one similar apicomedial
seta; palp article 2 with 4 apicalmedial marginal
spines and 4 submarginal setae. Plates of maxilla
2 extending equally, outer scarcely broader than
inner, outer with 4 apicolateral setae, inner with
4-6 medial setae fully to base. Inner plate of max-
illiped with 2 large thick apical spines, 3 apico-
facial setae, 6 medial setae; outer plate with 9
medial and apical spines, 6 apicolateral setae; palp
article 1 with one apicolateral seta, article 2 with
one apicolateral seta, medial margin of article 2
moderately to weakly setose, article 3 not protuber-
ant, with 3 facial setae, no lateral setae, nail of
article 4 long, with 2 accessory setules. Coxa 1
282 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
kRS
FIGURE 134.?Birubius gelarus, new species, holotype, female "n," 5.3 mm (/ ? juvenile female
"j," 4.7 mm; k ? female "k," 6.0 mm; Y ? eye).
scarcely expanded distally, anterior margin straight;
main ventral setae of coxae 1-4 = (11?13)?9?(8?10)?
(9-11), posteriormost seta of coxae 1-3 slightly
shortened; anterior and posterior margins of coxa
4 almost parallel, posterior margin convex, postero-
dorsal corner sharp, posterodorsal margin ordinary,
width-length ratio of coxa 4 = 15:16. Long pos-
terior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 2-(3-4)-(2-5)-(4-6), long ante-
riors = (2-4)-(10-ll)-0-0, short anteriors = 1-(1-
2)^-(2-3), no others. Gnathopods ordinary; width
ratios of articles 5-6 on gnathopods 1-2 = 29:35
and 27:31, length ratios = 64:67 and 58:60; pal-
mar humps ordinary, palms oblique; article 5 of
NUMBER 245 283
gnathopod 1 elongate, ovate, posterior margin
rounded-flat; article 5 of gnathopod 2 ovate, pos-
terior margin rounded. Pereopods 1-2 similar, fa-
cial setae formula on article 4 = 6 and 6, on article
5 = 6 and 6; main spine of article 5 extending to
M. 100+ on article 6, article 5 lacking proximo-
posterior spines; spine formula of article 6 = 4 + 4
and 3 + 4 plus middistal seta; acclivity on inner
margin of dactyls of pereopods 1-2 weak to obso-
lescent, emergent setule long, midfacial pluseta
ordinary. Coxae 5-7 posteroventral setule for-
mula = -3-4-1. Articles 4-5 of pereopods 3-4 of
ordinary width, facial spine rows sparse, facial ridge
formula on article 2 of pereopods 3-5 = 0-2-2,
article 2 of pereopod 3 with posterodorsal tooth;
width ratios of articles 2, 4, 5, 6 of pereopod 3 =
41:33:29:18, of pereopod 4 = 52:27:22:9, of pe-
reopod 5 = 76:21:19:7, length ratios of pereo-
pod 3 = 61:27:35:34, of pereopod 4 = 80:43:
45:50, of pereopod 5 = 100:26:21:21; article 2 of
pereopod 5 almost reaching middle of article 5,
ventral margin with long setae; medial apex of
article 6 finely combed, bearing 4 digital processes.
Posteroventral corner of epimeron 1 weakly pro-
tuberant sharply, posterior margin convex, with 2
setules, anteroventral margin with 4 medium setae,
posteroventral face with 2 medium setae; postero-
ventral corner of epimeron 2 with small sharp
tooth guarded by setule sinus, posterior margin
convex, with one additional setule, facial setae =
13, one middle pair set vertically; posteroventral
corner of epimeron 3 with large upturned tooth,
posterior margin almost straight, setose, with 5
long and short setules mostly near or in tooth
crotch, ventral margin with 6 setae mainly in mid-
dle of margin. Urosomite 1 with ventral setule at
base of uropod 1, articulation line incomplete,
short; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1-2 with articulate enlarged api-
cal nails, outer ramus of uropod 1 with 5-6 dorsal
spines, inner with 2, outer ramus of uropod 2 with
3?4 dorsal spines, inner lacking dorsomedial spine;
peduncle of uropod 1 with 2 apicolateral spines
and 5 basofacial setae, medially with 4-5 marginal
setae and spines, apicalmost an ordinary spine; pe-
duncle of uropod 2 with 6-7 dorsal spines, medially
with one small apical spine. Peduncle of uropod 3
with 5 ventral spines, dorsally with one lateral
spinule, 2 medial setules; rami submasculine, inner
extending to M. 85 on article 1 of outer ramus,
apex with one medium, one short setae, medial
margin with 4 setae, lateral margin with one, arti-
cle 2 of outer ramus short, 0.18, bearing 2 long setae,
medial margin of article 1 with 3 setae, lateral
margin with 4 acclivities, spine formula = long and
short, 2-2-2-2-2, setal formula = 0. Telson slightly
elongate, length-width ratio = 31:29, almost fully
cleft, each apex wide, rounded, lateral acclivity
deep, broad, bearing ordinary lateral setule, spine
next medial of length equal to setule, midlateral
setules subdiverse. Cuticle with ordinary bulbar
setules.
DESCRIPTION OF MALE (male "z," 5.16 mm).?
Following characters like B. gelarus, female maxilla
2, gnathopods, inner plate of maxilla 1 (long and
short setae), length of anterior setae on article 2
of pereopod 4; like B. babaneekus (16), right la-
cinia mobilis and pereopod 3, with possibility that
these parts are modified in males thereby making
identification difficult; antenna 1 with elongate
article 2 and primary flagellum, pereopod 5 with
short setae on article 2 and bulbars of cuticle with
long emergent setules, suggesting possibility that
this male is a distinctive species. Eyes hugely en-
larged, deeply stained burgundy. Article 1 on pe-
duncle of antenna 1 about 1.3 times as long as
wide, about 2.25 times as wide as article 2, ventral
margin with numerous setules, medial margin fuzzy,
produced dorsal apex with 4 setules; article 2
about 0.8 times as long as article 1, with ventral
cycle of 4 setae; primary flagellum with 10 articles,
about as long as peduncle, bearing several long
aesthetascs and calceoli on articles 1-6; accessory
flagellum with 6 articles, only half as long as pri-
mary flagellum. Spine formula of article 4 on an-
tenna 2 = 3-4-3, dorsomedial margins of articles
3^t densely fuzzy; facial spine formula of article
5 = 2-2, dorsal margin bearing 3 sets of male setae
and 2 calceoli, ventrodistal apex with 2 thin spines;
flagellum elongate, flagellar formula = (31-32), 2-5,
7, 9 . . . 31 or 1-4, 6, 8 . . . . Right lacinia mobilis
with distal branch forming weak blunt acclivity as
in B. babaneekus; left lacinia mobilis as in female
of B. gelarus; right rakers 4; left rakers 5 plus one
rudimentary; right molar with 6 spines, left molar
with 4 spines and one setule, no molar with plume;
palp article 2 with one short inner apical seta and
3 (2 in basal pair) other shorter inner setae, article
3 about 1.25 times as long as article 2, oblique apex
with 8 spine-setae, basofacial formula = 1 + 2-1 or
284 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
zGls
FIGURE 135.?Birubius gelarus, new species, holotype, female "n," 5.S mm (j ? juvenile "j," 4.7
mm; k =? female "k," 6.0 mm; z = male "z," 5.16 mm).
1 + 2-0. Inner plate of maxilla 1 bearing one long
apical pluseta, one shorter similar apicomedial seta,
2 apicolateral much shorter setae; palp article 2
with 4 apicalmedial marginal spines and 4 sub-
marginal setae. Inner plate of maxilla 2 with me-
dial setae to base. Inner plate of maxilliped with
5 medial setae, otherwise like B. babaneekus; palp
missing. Main ventral setae of coxae 1-4 = (11-12)-
9-8-10, coxae 1-3 with 1-2 additional anteroven-
tral setules besides normal member; coxa 4 like
figure of B. babaneekus but posterodorsal corner
rounded. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 2-3-5-5, long
anteriors = 4-5(+ 3 facial)-0-0, short anteriors =
1-1-3-4, no others. Gnathopods almost as thin as
female (see illustration). Pereopods 1-2, facial setae
NUMBER 245 285
formula on article 4 = 6-7 and 5; on article 5 =
5 and 6; spine formula of article 6 = 3 + 4 and
4 + 4 plus middistal seta; acclivity on inner margin
of dactyls of pereopods 1-2 weak, emergent setule
short, midfacial pluseta ordinary. Article 2 of pe-
reopod 3 subquadrate but not produced (see illus-
tration); pereopod 4 like female; pereopod 5 illus-
trated, article 2 of pereopod 5 with short ventral
setae, article 5 with enlarged posteroventral spine,
medial apex of article 6 finely combed, bearing 5
digital processes. Posteroventral corner of epimeron
1 rounded, weakly protuberant, posterior margin
deeply convex, corner with setule, and 3 other pos-
terior setules, anteroventral margin with 6 medium
setae, posteroverural face with 2 long setae; postero-
ventral corner of epimeron 2 rounded, weakly pro-
tuberant, with setule sinus above corner, posterior
margin weakly convex, with 2 setules, facial setae =
11; posteroventral corner of epimeron 3 with me-
dium tooth, posterior margin straight, with 2 setae
in crotch and 2 other posterior setules, ventral
margin with 7 setae; epimeron 3 with large seta
on posterodorsal margin set in weak notch. Outer
ramus of uropod 1 with 5 dorsal spines, inner with
2, outer ramus of uropod 2 with 3 dorsal spines,
inner with none; peduncle of uropod 1 with 2 api-
colateral spines and 4 basofacial setae, medially
with 5 thin marginal setae and spines plus apical
enlarged spine; peduncle of uropod 2 with 12 dor-
sal spines, medially with one apical spine. Peduncle
of uropod 3 with 5 ventral spines, dorsally with
one lateral spinule, 2 medial setules; rami mascu-
line, inner extending to M. 100+ on article 1 of
outer ramus, medial and lateral margins setose,
outer ramus lateral margin with 5 acclivities, spine
formula = 1-1?1?1?1?1, setal formula = 1-1-1-1-
1-1. Telson especially long, length-width ratio =
21:16, almost fully cleft, each apex bearing very
short lateral setule, spine next medial of length
equal to setule.
OBSERVATIONS.?Six out of 79 juveniles of this
species examined have a dorsomedial spine on the
inner ramus of uropod 2 but not necessarily on
both sides of the same individual.
ILLUSTRATIONS.?Following parts not illustrated,
substantially similar to those figured for B. baba-
neekus (16): right mandibular palp, inner plate of
maxilla 1, palp article 4 of maxilliped, coxae 1-4
(but setae fewer), pereopods 1-2; following parts
generally similar with exceptions noted in descrip-
tion: left mandibular palp article 2 (only one sub-
basal seta), outer plate and palp of maxilla 1 (but
outer plate with 11 spines), maxilliped. Illustrated
pereopod 4 taken from specimen other than that
used for illustrations of pereopods 3 and 5 but
magnification balanced.
HOLOTYPE.?NMV, female "n," 5.3 mm.
TYPE-LOCALITY.?CPBS Bl/1, 4 Aug 1964, West-
ern Port, Victoria, Australia, 7 m.
VOUCHER MATERIAL (all illustrated).?Type-local-
ity, female "k," 6.0 mm; CPBS 10E/3, juvenile fe-
male " j , " 4.7 mm; PPBES 919/4, male "z," 5.16 mm.
RELATIONSHIP.?This species differs from B.
babaneekus (16) conspicuously in the presence of
a dorsoposterior cusp on article 2 of pereopod 3
and in many minor characters, including broader
head and rostrum, better developed cusp on right
lacinia mobilis, smaller plume on molar, larger
main setae on inner plate of maxilla 1, presence of
medial setae basally on inner plate of maxilla 2,
narrower hands of gnathopods and longer setae on
article 4 of pereopod 4.
MATERIAL.?CPBS, 32 samples from 14 stations
(91); WPBES,3 samples from 3 stations (4); PPBES,
25 samples from 8 stations (106).
DISTRIBUTION.?Victoria: Western Port and Port
Philip Bay, 1-13 m, sand, silty sand, clayey sand,
Zostera.
18. Birubhis quearus, new species
FIGURE 136
DESCRIPTION OF MALE.?Head about 19 percent
of total body length, greatest width about 70 per-
cent of length; rostrum scarcely constricted, broad,
short, almost reaching middle of article 2 on an-
tenna 1. Eyes large, clear of pigment. Article 1 on
peduncle of antenna 1 about 1.2 times as long as
wide, about twice as wide as article 2, ventral mar-
gin with about 21 setules, weakly produced dorsal
apex with 4 setules; article 2 about 0.9 times as
long as article 1, with midventral cycle of 5 setae;
primary flagellum with 11 articles, about 0.95 times
as long as peduncle, bearing aesthetascs and one
calceolus each on articles 1-7; accessory flagellum
with 6 articles, only half as long as primary flagel-
lum. Spine formula on article 4 of antenna 2 =
3-4-3, dorsomedial margin of articles 3^1 bearing
fuzz, ventral margin with 5 groups of 1-2 long to
286 SMITHSONIAN CONTRIBUTION'S TO ZOOLOGY
FIGURE 136.?Birubius quearus, new species, holotype, male "b," 5.0 mm (/ ? juvenile "j,"
3.1 mm).
NUMBER 245 287
short setae, one ventrodistal long spine; article 5
about 1.1 times as long as article 4, facial spine
formula = 2-2, dorsal margin bearing 4 sets of
male setae (not visible in illustration) and 3 cal-
ceoli, ventral margin with 3 sets of 1-3 medium to
short setae, 2 thin ventrodistal long spines and one
setule; flagellum elongate, flagellar formula =
(31-32), 1-5, 7, 9 . . . 29 or 1-6, 8, 10 . . . 30. Man-
dibles with medium palpar hump; right incisor
with 3 teeth; left incisor with 4 humps in 2
branches; right lacinia mobilis bifid, very short and
broad, distal branch much shorter than proximal,
distal branch narrow, gaping, proximal branch
simple, pointed; left lacinia mobilis with 4 teeth,
middle teeth shortened; right rakers 3 plus one
rudimentary; left rakers 4 plus one rudimentary;
molar in form of elongate plaque, bulbous, each
molar with 4 primarily medium spines, none dis-
junct, no plume; palp article 1 short, article 2 with
one long inner apical seta and 2 other short inner
setae, article 3 about 1.25 times as long as article 2,
oblique apex with 8 spine-setae, basofacial for-
mula = 2-2. Inner plate of maxilla 1 ordinary,
bearing one medium simple pluseta, one similar
apicomedial seta, 2 apicolateral slightly shorter
setae; palp article 2 with 3 apicalmedial marginal
spines and 3 submarginal setae all highly distad,
palp elongate. Plates of maxilla 2 extending
equally, thin, of equal breadth, outer with 4 apico-
lateral setae, inner with 4 medial setae to base.
Inner plate of maxilliped with 2 large, thick apical
spines, 2 apicofacial setae, 3-4 medial setae; outer
plate with 9 medial and apical spines, 5 apico-
lateral setae; palp article 1 with one apicolateral
seta, article 2 with one apicolateral seta, medial
margin of article 2 moderately to weakly setose,
article 3 unprotuberant, with 2 facial setae, one
lateral seta, nail of article 4 medium, with 2 acces-
sory setules. Coxa 1 scarcely expanded distally, an-
terior margin almost straight; main ventral setae
of coxae 1-4 = 9-9-9-11, short, posteriormost seta
of coxae \~4 shortened; anterior and posterior mar-
gins of coxa 4 weakly divergent, posterior margin
oblique, straight, posterodorsal corner sharp, pos-
terodorsal margin short, straight, width-length
ratio of coxa 4 = 32:37. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
3-4-3-6, long anteriors = 4-5-1-0, short ante-
riors = 1-1-2-2, no others. Gnathopods ordinary;
width ratios of articles 5-6 on gnathopods 1-2 =
25:30 and 25:31, length ratios = 62:59 and 54:56;
palmar humps ordinary, palms oblique; article 5
of gnathopod 1 weakly elongate, ovate, posterior
margin rounded; article 5 of gnathopod 2 ovate,
posterior margin rounded, almost lobate. Pereopods
1-2 similar; facial setae formula on article 4 = 7
and 6, on article 5 = 5-6 and 6; main spine of arti-
cle 5 extending to M. 100 on article 6, article 5
lacking proximoposterior spines; spine formula of
article 6 = 3 + 4 and 3 + 4 plus middistal seta;
acclivity on inner margin of dactyls of pereopods
1-2 weak, emergent setule long, midfacial pluseta
ordinary. Coxae 5-7 posteroventral setule for-
mula = 4-5-1. Articles 4-5 of pereopods 3-4 of
ordinary width, facial spine rows moderately de-
veloped, facial ridge formula on article 2 of pereo-
pods 3-5 = 0-2-2; width ratios of articles 2, 4, 5,
6 of pereopod 3=40:30:32:16, of pereopod 4 =
56:28:23:9, of pereopod 5=78:20:19:5, length
ratios of pereopod 3 = 62:25:27:33, of pereopod
4 = 78:45:44:45, of pereopod 5 = 104:27:23:18;
article 2 of pereopod 5 exceeding apex of article
4, ventral setules large and numerous; article 5
with enlarged posteroventral spine; medial apex of
article 6 smooth, bearing 2 digital processes. Pos-
teroventral corner of epimeron 1 rounded, posterior
margin deeply convex, with 2 setules, anteroventral
margin with 6-7 medium setae, posteroventral face
with 2-3 long setae; posteroventral corner of epi-
meron 2 rounded-quadrate, posterior margin con-
vex, with 2-3 setules, facial setae = 12-13, occa-
sional pairs set vertically or not; posteroventral
corner of epimeron 3 with medium tooth, posterior
margin weakly convex, scalloped, with 4 setule
notches, ventral margin with 6-7 setae mainly ante-
rior, occasional pair set vertically; epimera 1-3
with setule on posterodorsal margin set in weak
notch. Urosomite 1 with 2 ventral setules at base
of uropod 1, articulation line complete; urosomite
3 protuberant dorsally. Rami of uropods 1-2 with
articulate enlarged long sharp apical nails, outer
ramus of uropod 1 with 5-6 dorsal spines, inner
with 2, outer ramus of uropod 2 with 4 dorsal
spines, inner with none; peduncle of uropod 1 with
3 apicolateral spines and 6 basofacial setae, medi-
ally with 5 marginal spines plus apical enlarged
spine; peduncle of uropod 2 with 13 dorsal spines,
medially with one small apical spine. Peduncle of
uropod 3 with 2 ventral spines, dorsally with one
lateral spine, one medial setule; rami masculine,
288 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
inner extending to M. 100+ on article 1 of outer
ramus, apex with 3 setae, medial and lateral mar-
gins setose, article 2 of outer ramus short, 0.12,
bearing 2 long setae, apicomedial margin of article
1 setose, lateral margin with 5 acclivities, spine for-
mula = 0-0-1-1-1-1, setal formula = 1-1-1-1-1-1.
Telson long, length-width ratio = 7:6, almost fully
cleft, each apex wide, rounded, lateral acclivity nar-
row, weak, bearing short lateral setule, spine next
medial of length equal to setule, each lobe with long
dorsal row of denticles, midlateral setules diverse.
Cuticle with ordinary bulbar setules surrounded by
plaques forming clear spaces on surface, bearing fine
striations in form of linear fingerprint pattern, emer-
gent setules plumose.
DESCRIPTION OF JUVENILE (juvenile " j , " 3.1 mm).
?Female facies. Article 2 of antenna 1 shorter
than in male, with 4 ventral setae; primary flagel-
lum 7-articulate; accessory flagellum 4-articulate.
Article 4 of antenna 2 with facial spine formula of
1-3-4-2, of article 5 = 2-2, distal apex with 3 long
to medium spines, flagellum 7-articulate. Mandibles
as in male, basofacial setal formula on article 3 =
0-1, apex with 6 spine-setae. Long setal formula on
coxae 1-4 = 6-6-6-7; posterodistal corner of coxa
4 rounded. Article 5 of gnathopod 1 slightly more
elongate than in male. Pereopod 5 with 8 elongate
ventral setae on article 2, no enlarged spine on
article 5. Setal formulas on epimera, anteroventral
epimeron 1 = 2 , posteroventral = 1, posterior =
1-2 setules, posteroventral corner with small sharp
tooth, epimeron 2 facial = 6-7, posteroventral = 1,
epimeron 3 ventral = 2, posterior = 2 long setules,
epimeron 1 with posterodorsal setule, epimera 2-3
with long posterodorsal seta. Spine formulas on
uropods, peduncle of uropod 1 = 2 , of uropod
2 = 4, outer rami of uropod 1 = 3, of uropod 2 = 2 ,
inner rami of uropod 1 = 1, of uropod 2 = 0, ven-
tral of uropod 3 peduncle = 4, basofacial setae of
uropod 1 = 4 , spine formula on outer ramus of
uropod 3 = 1-2-2 (elongate), no setae. Apical spines
of telson elongate but setules also elongate. Eyes
not apparent, probably destroyed in preservative.
OBSERVATIONS.?The juvenile adds only a few
points to the probable female morphology: the
elongate ventral setae on article 2 of pereopod 5
and the posteroventral points on epimera 1-2;
otherwise the female is suspected to have the stand-
ard female morphology of eyes, epimera, uropod 3.
ILLUSTRATIONS.?Following unfigured parts like
those of B. babaneekus (16): antenna 1 but article
2 more elongate and with more ventral setules on
article 1, maxillae 1-2, maxilliped except article 4
as described, coxae, (except setae much shorter),
gnathopods, pereopods 1-2, pereopod 3 (except as
partially illustrated). Following parts like B. gela-
rus (17): upper lip, pereopod 4. Dorsal male setae
on article 5 of antenna 2 not shown (rotated be-
hind); male epimeron 3 showing 7 ventral setal
holes, eighth hole set vertically above showing posi-
tion of seta 7 on right epimeron, latter bearing 6
ventral setae and one disjunct seta; pereopod S
enlarged 1.1 times over pereopod 5.
HOLOTYPE.?NMV, male "b," 5.0 mm.
TYPE-LOCALITY?CPBS A4(T)/1. 10 Oct 1964,
Western Port, Victoria, Australia, 9 m.
VOUCHER MATERIAL.?CPBS 51N/1, juvenile " j , "
3.1 mm (illus.).
RELATIONSHIP.?This species differs from either
or both B. babaneekus (16) and B. gelarus (17) in
the much smaller tooth of epimeron 3, the large
setae on the inner plate of maxilla I and the pres-
ence of only 4 spines on the molars.
MATERIAL.?CPBS, 2 samples from 2 stations (2).
DISTRIBUTION.?Victoria, Western Port, 9-14 m.
19. Birubius narus, new species
FIGURES 137, 138
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total body length, greatest width about 60
percent of length; rostrum strongly constricted,
narrow, reaching middle of article 2 on antenna 1.
Eyes medium to large, largely occluded with pig-
ment, ommatidia especially large. Article 1 on pe-
duncle of antenna 1 about 1.3 times as long as
wide, about 1.75 times as wide as article 2, ventral
margin with about 11 setules, weakly produced
dorsal apex with one setule; article 2 about 0.73
times as long as article 1, with ventral crescent of
5 setae; primary flagellum with 10 articles, about
as long as peduncle, bearing aesthetascs; accessory
flagellum with 8 articles. Spine formula on article
4 of antenna 2 = 1-2-4-3, dorsal margin with
notch bearing one seta and one spine, ventral mar-
gin with 3 groups of 2-4 long to medium setae,
one ventrodistal long spine; article 5 about 0.7
times as long as article 4, facial spine formula =
1-2, dorsal margin naked, ventral margin with 2
NUMBER 245
289
s
FIGURE 137.?Birubius narus, new species, holotype, female "a," 2.90 mm.
sets of 2-3 long to medium setae, 2 ventrodistal
long to medium spines; flagellum about 1.35 times
as long as articles 4-5 of peduncle combined, with
10 articles. Mandibles with medium palpar hump;
right incisor with 3 teeth; [left incisor and lacinia
mobilis damaged, not analyzed]; right lacinia mo-
bilis simple, denticulate; right rakers 8; left rakers
8 plus one rudimentary; molar in form of elongate
plaque, right molar with 7 primarily long spines,
none disjunct, left molar with 5 primarily long
spines, none disjunct, each molar with plume; palp
article 1 short, article 2 with one long inner apical
290 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
seta and one other shorter inner seta, article 3
about 0.9 times as long as article 2, oblique apex
with 6 spine-setae, basofacial formula = 0. Inner
plate of maxilla 1 ordinary, bearing one long api-
cal pluseta, one shorter apicomedial seta, 2 apico-
lateral shorter setae; palp article 2 with 2 apical-
medial marginal spines and 4 marginal setae, one
of latter apicolateral. Plates of maxilla 2 extending
subequally, of subequal width, outer with one api-
colateral seta, inner with one medial seta. Inner
plate of maxilliped with one large thick apical
spine, 2 apicofacial setae, 3 medial setae; outer
plate with 4 medial and apical spines, no apico-
lateral setae; palp article 2 with one apicolateral
seta, medial margin of article 2 moderately setose,
article 3 unprotuberant, with 3 facial setae, no
lateral setae, nail of article 4 long, with 2 accessory
setules. Coxa 1 strongly expanded distally, anterior
margin concave; main ventral setae of coxae 1-4 =
2-2-2-0, posteriormost seta of coxae 1-3 shortened;
anterior and posterior margins of coxa 4 weakly
divergent, posterior margin very convex, postero-
dorsal corner rounded, posterodorsal margin short,
concave, width-length ratio of coxa 4 = 37:41.
Long posterior setae on article 2 of gnathopods
1-2 and pereopods 1-2 ? 2-2-1-, long anteriors ?
3-3-0-1, short anteriors = (l-2)-4-3-2, no others.
Gnathopods ordinary, hands stout; width ratios
of articles 5-6 on gnathopods 1-2 = 23:40 and
24:41, length ratios = 68:65 and 63:60; palmar
humps ordinary, palms weakly oblique; article of
5 of gnathopod 1 elongate, ovate, posterior margin
flat; article 5 of gnathopod 2 elongate, ovate, pos-
terior margin rounded. Pereopod 2 slightly stouter
than pereopod 1, especially article 4; facial setae
formula on article 4 = 2 and 2, on article 5 = 2
and 2; main spine of article 5 extending to M. 75
on article 6, article 5 with one proximoposterior
spine, spine formula of article 6 = 3 + 4 and 4 + 4
plus middistal seta, one spine especially long; ac-
clivity on inner margin of dactyls of pereopods 1-2
blunt, produced as tooth, emergent setule short,
midfacial pluseta ordinary. Coxae 5-7 posteroven-
tral setule formula = 1-1-1. Articles 4-5 of pereo-
pods 3^4 broad, facial spine rows moderately de-
veloped, facial ridge formula on article 2 of pereo-
pods 3-5 = 0-2-2, anterior ridge on pereopod 5
short, width ratios of articles 2, 4, 5, 6 of pereopod
3 = 46:42:36:15, of pereopod 4 = 61:42:29:13, of
pereopod 5 = 78:17:16:8, length ratios of pereo-
pod 3 = 70:26:35:37, of pereopod 4 = 78:46:
32:43, of pereopod 5=101:23:21:25: article 2
of pereopod 5 reaching middle of article 5; me-
dial apex of article 6 finely combed, bearing obso-
lescent digital processes. Posteroventral corner of
epimeron 1 rounded-quadrate, posterior margin
weakly convex, anteroventral margin with one me-
dium seta, posteroventral face with 2 long to short
setae set vertically; posteroventral corner of epi-
meron 2 rounded-quadrate, weakly protuberant,
posterior margin undulant, facial setae = 5, poste-
riormost pair set vertically; posteroventral corner
of epimeron 3 with medium to large tooth, poste-
rior margin weakly convex, with 2 setae in crotch;
epimera 1-3 with seta on posterodorsal margin set
in weak notch. Urosomite 1 with 2 midventral
setules, articulation line incomplete, short; uroso-
mite 3 weakly protuberant dorsally. Rami of uro-
pods 1-2 with articulate enlarged apical nails, outer
ram us of uropod I with 4 dorsal spines, inner with
2, outer ramus of uropod 2 with 3 dorsal spines,
inner with one dorsomedial spine; peduncle of
uropod 1 with one apicolateral spine and one baso-
facial seta, medially with 2 marginal spines, apical-
most ordinary; peduncle of uropod 2 with 4 dorsal
spines, medially with one apical spine. Peduncle
of uropod 3 with 3 ventral spines, dorsally with
one lateral spine, one medial setule; rami feminine,
inner extending to M. 75 on article 1 of outer
ramus, apex with one long seta, medial and lateral
margins naked, article 2 of outer ramus elongate,
0.28, bearing 2 medium to long setae, medial mar-
gin of article 1 with one seta, lateral margin with
3 acclivities, spine formula = 1-2-2-2, setal for-
mul = 0. Telson ordinary, length-width ratio =
1:1, not fully cleft, each apex of medium breadth,
rounded, lateral acclivity weak, bearing long lat-
eral setule, spine next medial slightly shorter than
setule, midlateral setules diverse. Cuticle with ordi-
nary bulbar setules, surface bearing fine striations
in form of linear fingerprint pattern.
ILLUSTRATIONS.?Three lateral spines on article 6
of pereopod 1 blackened so as to distinguish them
from medial row; head distorted dorsally; prebuc-
cal lateral view not ascertained, damaged.
HOLOTYPE.?AM, female "a," 2.90 mm. Unique.
TYPE-LOCALITY.?EBS 20, 25 Apr 1972, Jervis
Bay, Murray's Beach, New South Wales, Australia,
6 m, Zostera.
RELATIONSHIP.?This species bears superficial re-
NUMBER 245 291
FIGURE 138.?Birubius narus, new species, holotype, female "a," 2-90 mm.
292 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
semblance to the B. babaneekus group (16-18) be-
cause of the large tooth on epimeron 3, the absence
of accessory nails on uropod 1, and the simple right
lacinia mobilis but in other respects differs widely
from that group and is assumed to have no rela-
tionship to it. Some of the characters distinguishing
B. narus from the B. babaneekus group are: the
denticulations found on the right lacinia mobilis
and neighboring raker spines; broad expansion of
coxa 1 and reduction of anterior coxal setae; ab-
sence of setae on coxa 4; poorly developed baso-
facial setae on uropod 1; crowded setae of epi-
meron 2; narrow rostrum; elongate article 2 on
outer ramus of uropod 3; broader articles of pe-
reopods 3-4; small size of the outer plate on the
maxillipeds; absence of conspicuous ventral setae
on article 2 of pereopod 5; elongate article 5 of
gnathopod 2; expanded hands of gnathopods;
strong dactylar acclivity on pereopods 1-2; and
the shorter main spine on article 5 of pereopods
1-2.
Birubius narus differs from B. panamunus (1) in
the absence of facial setae on epimeron 3 and the
absence of long setae on coxa 4, but is closer to
B. apart (6) in that this latter species also lacks
setae on coxa 4.
Birubius narus is a conceivable ancestral type to
the B. taldeus (34-37) group because of resem-
blances in rostrum and coxa 1 among numerous
other details, but differs from the B. taldeus group
in the large tooth of epimeron 3 and the absence
of accessory apical setules on the rami of uropods
1-2. In the latter character B. narus is closer to
B. eake (32), a satellite member of the B. taldeus
group.
MATERIAL.?EBS, one sample (1).
DISTRIBUTION.?New South Wales, Jervis Bay,
Murray's Beach, 6 m, Zostera.
20. Birubius gambodeni, new species
FIGURES 139-143
DESCRIPTION OF FEMALE (holotype, female "c,"
10.6 mm, from Western Australia).?Head about
16 percent of total body length, greatest width
about 70 percent of length; rostrum scarcely con-
stricted, narrow apically, exceeding middle of arti-
cle 2 on antenna 1. Eyes medium, largely occluded
with pigment. Article 1 on peduncle of antenna 1
about 1.4 times as long as wide, about 2.5 times
as wide as article 2, ventral margin with numerous
setules, unproduced dorsal apex with 2 setules;
article 2 about 0.65 times as long as article 1, with
ventral cycle of 5 setae; primary flagellum with
13-14 articles, about 0.66 times as long as peduncle,
bearing several short aesthetascs; accessory flagel-
lum with 12 articles. Spine formula on article 4
of antenna 2 = 1-3-4-7. dorsal margin with 2
notches bearing 9 setae and spines, ventral margin
with about 9 groups of 2 long to medium setae,
one ventrodistal medium spine; article 5 about 0.75
times as long as article 4, facial spine formula =
3-3, dorsal margin with 2-3 setules, ventral margin
with 5 sets of 2 long to short setae, 2 ventrodistal
long to medium spines: flagellum as long as articles
4-5 of peduncle combined, with 15 articles. Man-
dibles with strong palpar hump; right incisor with
3 teeth; left incisor with 3 humps in 2 branches;
right lacinia mobilis bifid, distal branch shorter
than proximal, broad, subfalrate, proximal branch
simple, pointed, with marginal denticles; left la-
cinia mobilis with 4 teeth; right rakers 9 plus 1-2
rudimentaries; left rakers 10 plus one rudimentary;
molar in form of elongate plaque, right molar with
8 primarily long spines plus one short spine strongly
disjunct, left molar with 7 primarily long spines,
plus one short spine strongly disjunct; palp article
1 short, article 2 with 3 long inner apical setae and
2-3 other shorter inner setae plus 2-3 outer setae,
article 3 about 1.1 times as long as article 2, oblique
apex with 11 spine-setae, basofacial formula = 2-1
or 1-1. Inner plate of maxilla 1 ordinary, bearing
one long apical pluseta, one similar apicomedial
seta, 2 apicolateral much shorter setae; palp article
2 with 3 apicalmedial marginal spines and 4 sub-
marginal setae. Plates of maxilla 2 extending sub-
equally, of equal width, outer with 6 apicolateral
setae, inner with 3 medial setae. Inner plate of
maxilliped with one large thick apical spine, 3
apicofacial setae, 4 medial setae; outer plate with
7 medial and apical spines, 2 apicolateral setae;
palp articles 1-2 with several lateral setae, medial
margin of article 2 strongly setose, article 3 pro-
tuberant, with 5 facial setae, group of 4 lateral
setae, nail of article 4 medium, with 2 accessory
setules. Coxa 1 expanded distally, anterior margin
almost straight; main ventral setae of coxae 1?4 ?
8-10-10-0, posteriormost seta of coxae 1-3 short-
ened; anterior and posterior margins of coxa 4
NUMBER 245 293
s
FIGURE 139.?Birubius gambodeni, new species, holotype, female "c," 10.6 mm.
294 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
weakly divergent, posterior margin almost straight,
posterodorsal corner rounded, posterodorsal margin
undulant, width-length ratio of coxa 4 = 17:20.
Long posterior setae on article 2 of gnathopods
1-2 and pereopods 1-2 = (5-8)-8-(7-8)-10, long
anteriors = 6-18-0-0, short anteriors = 5-10-7-4.
Gnathopods ordinary; width ratios of articles 5-6
on gnathopods 1-2 = 29:31 and 29:31, length
ratios = 65:65 and 60:65; palmar humps ordi-
nary, palms oblique; article 5 of gnathopod 1
slightly elongate, ovate, posterior margin flat; arti-
cle 5 of gnathopod 2 ovate, posterior margin
rounded. Pereopods 1-2 similar, facial setae for-
mula on article 4 = 8 and 7, on article 5 = 8 and 8;
main spine of article 5 extending to M. 100 on
article 6, article 5 lacking proximoposterior spines;
spine formula of article 6 = 5 + 6 plus middistal
seta; acclivity on inner margin of dactyls of pereo-
pods 1-2 weak, emergent setule short, midfacial
pluseta ordinary, highly distad. Coxae 5-7 postero-
ventral seta formula = (11?12)?5?4. Articles 4-5 of
pereopods 3-4 broad to medium, facial spine rows
moderately developed, facial ridge formula on arti-
cle 2 of pereopods 3-5 = 0-2-2, anterior on pereo-
pod 5 short; width ratios of articles 2, 4, 5, 6
of pereopod 3 = 43:45:35:15, of pereopod 4 =
70:39:27:12, of pereopod 5 = 80:25:22:8, length
ratios of pereopod 3 = 78:28:37:40, of pereopod
4 = 90:50:43:55, of pereopod 5 = 100:30:25:28;
article 2 of pereopod 5 reaching apex of article
4; medial apex of article 6 finely combed,
bearing 6-7 digital processes. Posteroventral cor-
ner of epimeron 1 subquadrate, posterior mar-
gin weakly convex, with 2 setules above corner,
anteroventral margin with 9 short to medium setae,
posteroventral face with 3 medium setae; postero-
ventral corner of epimeron 2 weakly protuberant,
posterior margin straight, weakly undulant, with
setule and 4 setae, facial setae = 1 1 , three pairs set
vertically, face dorsally and posterior to ridge with
3 setae; corner of epimeron 3 with small tooth,
posterior margin almost straight, strongly setose,
ventral margin naked, face with oblique irregular
row of 6 setae nearly extending to middle. Uroso-
mite 1 naked, articulation line short; urosomite 3
weakly protuberant dorsally. Rami of uropods 1-2
with articulate enlarged apical nails, outer ramus
of uropod 1 with 10 dorsal spines, inner with 3,
outer ramus of uropod 2 with 9 dorsal spines, inner
with 2 dorsomedial spines; peduncle of uropod 1
with 6 apicolateral spines and 5 basofacial setae,
medially with many marginal setae plus apical en-
larged spine; peduncle of uropod 2 with 14 dorsal
spines, medially with one small apical spine. Pe-
duncle of uropod 3 with 6 ventral spines, dorsally
with one lateral spine, one medial spine; rami sub-
masculine, inner extending to M. 87 on article 1
of outer ramus, apex with 2 setae, medial margin
setose, article 2 of outer ramus short, 0.12, bearing
2 medium to long setae, apicomedial margin of
article 1 setose, lateral margin with 5 acclivities,
spine formula = 2-2-2-2-2-2. setal formula = 0.
Telson ordinary, length-width ratio = 15:14, al-
most fully cleft, each apex wide, rounded, often
with 2 midapical setae, lateral acclivity deep, bear-
ing short lateral setule, spine next medial longer
than setule. midlateral setules weakly diverse. Cu-
ticle with bulbar setules mixed with pipes bearing
apical horse*ollar, emergent setules especially short.
OBSERVATIONS.?Pereopods 1-2 of holotype each
with 7 and 4 medium to short anterior setae; pre-
buccal complex convex anteriorly from lateral view,
ventrally truncate or weakly excavate with mid-
hump from anterior view, articulation damaged
during dissection.
VARIATIONS.?Female "h," 11.8 mm, PPBES
903/1, from Victoria: Disjunct spine on molars
very thin, simple; distal branch of right lacinia
mobilis perfectly simple, unexpanded; hands of
gnathopods stouter than in Western Australian
holotype; apical nail on dactyl of pereopods 1-2
deeply submerged. This specimen is illustrated to
show minor differences in other characters, mainly
setal and spinal counts. Right mandible with 8 +
one rudimentary raker spines, left mandible with
10 + one rudimentary, note accessory teeth on first
2 raker spines of left mandible (illustrated), note
fork in large clear spine on both mandibular mo-
lars; in illustration of mandibular molars truncate
spines broken apically; primary flagellum and ac-
cessory flagellum of antenna 1 each with 12 articles,
article 2 of peduncle on antenna 1 with 10 ventral
setae in main group, article 1 with 19 feathered
setules posteriorly; antenna 2 with 14 articles in
flagellum; left mandibular palp article 3 with
3 + 2 facial (basal) setae, right palp with 3 + 1
(illustrated); article 2 of gnathopod 1 with 5 pos-
terior setae (3 at M. 33, 2 at M. 66), gnathopod
2 with 7 or 8 posterior setae; pereopods 1-2 lack-
ing proximoposterior spine on article 5.
NUMBER 245 295
R3
FIGURE 140.?Birubius gambodeni, new species, holotype, female "c," 10.6 mm.
296 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 141.?Birubius gambodeni, new species, female "h," 11.8 mm.
NUMBER 245 297
FIGURE 142-?Birubius gambodeni, new species, female "h," 11.8 mm.
Other Victorian specimens with left molarial dis-
junct spine thin, right spine as in holotype, right
lacinia mobilis as in holotype.
ILLUSTRATIONS.?Spines on right mandibular
molar of holotype not flattened.
HOLOTYPE.?WAM, female "c," 10.6 mm.
TYPE-LOCALITY.?JLB AUS 12, 30 Sep 1968,
Middleton Beach, Albany, Western Australia, in-
tertidal wash of sandy rocks, coralline algae.
VOUCHER MATERIAL.?PPBES 903/1, female "h,"
11.8 mm (illus.). Male unknown.
RELATIONSHIP.?Birubius gambodeni forms the
model for a group of species, 20-27, characterized
by strongly bifid right lacinia mobilis, each branch
of which is simple or nearly so, and by uropods 1-2
lacking accessory nails. All species lack long setae
on coxa 4 but species 21-27, in contrast to B. gam-
bodeni, bear one or more posteroproximal spines
298 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 143.?Birubius gambodeni, new species, female "h," 11.8 mm.
on article 5 of pereopods 1-2.
Birubius gambodeni is placed in both keys A and
B of Birubius because the distal branch of the right
lacinia mobilis is intermediate between simple and
complex.
A strong relationship is seen between B. gam-
bodeni and the B. myallus group (4-5) because of
correspondence between facial setae of epimeron
3, absence of long setae on coxa 4, slightly con-
stricted rostrum, similar spination and setation pat-
terns on the peduncles of uropods 1-2 and coxae
1-3, and because certain specimens of B. myallus
show a reduction in the distal branch of the right
lacinia mobilis. Birubius gambodeni differs from
the B. myallus group in the absence of accessory
nails on uropods 1-2.
In the B. gambodeni group specific distinctions
can be made in 4 characters: (1) short or long arti-
cle 5 of gnathopod 1; (2) long or short anterior
facial ridge on article 2 of pereopod 5; (3) dis-
persed or crowded facial setae on epimeron 2; (4)
elongate or shortened article 2 on the outer ramus
of uropod 3. The 8 species of the group can be
divided into 5 groups on this basis. The first alter-
native in each category is selected as primitive. No
species has fewer than 2 of the characters in non-
primitive condition so that numerous more primi-
tive combinations can be hypothesized than have
actually been found, indicating a wide divergence
of each group from the hypothetically primitive
state. Group 1, composed of B. gambodeni, B. maa-
mus (21), and B. kokorus (25), has characters 2 and
4 in advanced state; group 2, composed of B. mung-
gai (27) has characters 1, 3, 4 in advanced state;
group 3, composed of B. lowannus (22) and B.
kyeemus (23) has characters 1, 2, and 3 in advanced
state; group 4, composed of B. batei (24), has char-
acters 1 and 4 in advanced state, and B. kinkus (26)
has all characters in advanced state.
In Figure 82 the 8 species are placed in a slightly
different arrangement reflecting a combination of
characters concerning epimeron 3, the elongate
wrist of the gnathopods and basofacial setation on
uropod 1. Three kinds of epimeral setation are
considered significant: facially scattered, facially or-
ganized, and posteriorly aligned. In this way B.
batei is brought into the ancestral position of B.
jirrandus (31).
MATERIAL.?PPBES, 5 samples from 4 stations
(9); JLB AUS, one sample (1).
NUMBER 245 299
DISTRIBUTION.?Victoria, Port Phillip Bay, 5-7 m,
sand, sandy silt; Western Australia, Albany, 0 m.
21. Birubius maamus, new species
FIGURES 144-146
DESCRIPTION OF FEMALE.?Head about 17 per-
cent of total body length, greatest width about 60
percent of length; rostrum constricted, narrow,
elongate, reaching middle of article 2 on antenna 1.
Eyes small to medium, largely occluded with pig-
ment. Article 1 of peduncle on antenna 1 almost
1.5 times as long as wide, about twice as wide as
article 2, ventral margin with about 10-12 setules,
weakly produced dorsal apex with 2 setules; article
2 about 0.7 times as long as article 1, with proximo-
ventral cycle of 4 setae; primary flagellum with
9-11 articles, about 0.7 times as long as peduncle,
bearing several long aesthetascs distally; accessory
flagellum with 7-9 articles. Spine formula on article
4 of antenna 2 - 1-3^-3 or 1-3-4-4 or 1-3-5-3,
dorsal margin with notch bearing 3 setal-spines,
ventral margin with 6 groups of 2 long and me-
dium setae, one ventrodistal long spine; article 5
about 0.7 times as long as article 4, facial spine
formula = 1-3, dorsal margin naked, ventral mar-
gin with 4 sets of 2 long and short setae, 2 ventro-
distal medium spines; flagellum about 0.8-0.9
times as long as articles 4-5 of peduncle combined,
with 10-12 articles. Mandibles with medium palpar
hump; right incisor with 3 teeth; left incisor with
4 teeth in 2 branches; right lacinia mobilis bifid,
distal branch shorter than proximal, distal branch
simple, proximal branch simple, pointed, with mar-
ginal denticles; left lacinia mobilis with 4 teeth;
right rakers 7-9 plus 1-2 rudimentaries, left rakers
9-10 plus 1-2 rudimentaries; molar in form of
hump demarcated mainly by spines, right molar
with 6-8 primarily long spines, none disjunct, left
molar with 7-8 primarily long spines, none dis-
junct, only left molar with plume, palp article 1
short, article 2 with one short inner apical seta and
1-3 other shorter inner setae, article 3 about 0.9-1.1
times as long as article 2, oblique apex with 10-14
spine-setae, basofacial formula = 0-1. Inner plate
of maxilla 1 large, bearing one long apical pluseta,
one similar or shorter apicomedial seta, 2 apico-
lateral much shorter setae (one hidden in illustra-
tion); palp article 2 with 3 apicalmedial marginal
spines and 4-5 submarginal setae. Plates of maxilla 2
extending equally, outer scarely broader than inner,
with 4-5 apicolateral setae, inner with 1-2 medial
setae. Inner plate of maxilliped with one large, thick
apical spine, 2 apicofacial setae, 4 medial setae;
outer plate with 6 medial and apical spines, 2-3
apicolateral setae; palp article 2 with 2 groups of
2-A apicolateral setae, medial margin of article 2
strongy setose, article 3 protuberant, with 4 facial
setae, 1-3 lateral setae, nail of article 4 short, with
2 accessory setules. Coxa 1 weakly expanded dis-
tally, anterior margin weakly convex; main ventral
setae of coxae 1^ ? = 11-9-10-0, posteriormost seta
of coxae l-2(or 3) slightly shortened; anterior and
posterior margins of coxa 4 almost parallel, poste-
rior margin convex, posterodorsal corner rounded,
posterodorsal margin short, concave, width-
length ratio of coxa 4 = 7:8. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 6-(4-5)-(4-5)-(4-5), long anteriors = 0- (2-
3)-0-0, others not counted. Gnathopods generally
ordinary; width ratios of articles 5-6 on gnathopods
1-2 = 27:32 and 27:35, length ratios = 57:65
and 51:62; palmar humps ordinary, palms oblique;
article 5 of gnathopod 1 ovate, posterior margin
flat; article 5 of gnathopod 2 ovate, posterior mar-
gin rounded. Pereopods 1-2 similar, facial setae
formula on article 4 = (5-6), on article 5 = (6-7);
main spine of article 5 extending to M. 95 on arti-
cle 6, article 5 with one proximoposterior spine;
spine formula of article 6 = 5 + 5 plus middistal
seta; acclivity on inner margin of dactyls of pereo-
pods 1-2 weak, emergent setule medium, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule
formula = (4-5)-(4-5)-(4-6). Articles 4-5 of pereo-
pods 3-4 broad, facial spine rows moderately to
well developed, facial ridge formula on article 2 of
pereopods 3-5 = 0-2-2, anterior ridge on pereopod
5 short, width ratios of articles 2, 4, 5, 6 of pe-
reopod 3=43:43:36:17, of pereopod 4 = 68:42:
31:13, of pereopod 5 = 85:24:20:8, length ratios
of pereopod 3 = 72:31:36:40, of pereopod 4 =
80:50:40:50, of pereopod 5 = 107:30:26:27; article
2 of pereopod 5 reaching middle of article 5;
medial apex of article 6 finely combed, bearing 5-6
digital processes. Posteroventral corner of epimeron
1 weakly protrusive, posterior margin weakly con-
vex, with 2 setules, anteroventral margin with 6-7
short to medium setae, posteroventral face with 2-3
long setae; posteroventral corner of epimeron 2
300 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 144.?Birubius maamus, new species, holotype, female "w," 6.7 mm (a ?= male "a," 5.6
mm; n ? female "n," 6.6 mm; u ? male "u," 6.1 mm; y ? female "y," 7.9 mm; Y ? left
lacinia mobilis).
NUMBER 245
uGlt
FIGURE 145.?Birubius maamus, new species, holotype, female "w," 6.7 mm (a = male "a," 5.6
mm; n ?= female "n," 6.6 mm; u = male "u," 6.1 mm; y = female "y," 7.9 mm).
302 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
weakly protuberant, guarded by setule sinus, poste-
rior margin weakly undulant, facial setae = 6 in
horizontal row and 2-A irregularly disposed poste-
rior setae; posteroventral corner of epimeron 3 with
setule sinus and small tooth, posterior margin
straight, with 3^1 setule notches and 2 long setae
near tooth, ventral margin with 2 posterior setae
near tooth, face with oblique row of 4 setae and 2
others or 6 scattered setae, epimeron 3 with setule on
posterodorsal margin set in weak notch. Urosomite 1
with ventral setule at base of uropod 1, articulation
line short; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1?2 with articulate enlarged
apical nails, rami of uropods 1-2 lacking accessory
nails, outer ramus of uropod 1 with 6-8 dorsal
spines, inner with 2-3, outer ramus of uropod 2
with 5-6 dorsal spines, inner with 1-2 dorsomedial
spines; peduncle of uropod 1 with 4 apicolatcral
spines and 4-5 basofacial setae, medially with 4-7
marginal spines, apicalmost enlarged; peduncle of
uropod 2 with 12 dorsal spines, with one small
apical spine medially. Peduncle of uropod 3 with
6-7 ventral spines, dorsally with one lateral spine,
one medial spine; rami feminine, inner extending
to M. 68 on article 1 of outer ramus, apex with one
seta, medial and lateral margins naked, article 2 of
outer ramus short, 0.17, bearing 2 long setae, apico-
medial corner of article 1 naked, medial margin
with 2 setae, lateral margin with 4 acclivities, spine
formula = 2-2-2-2-2 setal formula = 0. Telson
ordinary, length-width ratio = 12:11, not fully
cleft, each apex wide, rounded, lateral acclivity
weak, bearing ordinary lateral setule, spine next
medial of length subequal to setule, midlateral
setules diverse. Cuticle with ordinary bulbar setules,
emergent setules ordinary.
DESCRIPTION OF MALE.?Rostrum slightly longer
than in female. Article 2 of antenna 1 with 4 ven-
tral setae; primary flagellum with 9 articles, one
calceolus each on articles 1-4; accessory flagellum
with 7-8 articles. Facial spine formula on article 4
of antenna 2 = 2-4-3 or 2-4r-2, on article 5 = 3 or
2, article 5 with 2-3 dorsal sets of male setae and 2
large calceoli (not visible in illustration), ventro-
distal apex with 2 thin spines and one setule, fla-
gellar formula = (29-33), 2, 4, 6 . . . penultimate
or 2-A, 6, 8 . . . or 2, 3, 5, 7 . . . . Proximal branch of
right lacinia mobilis slender; mandibular rakers
right = 7-8 plus one rudimentary, left = 6-7 plus
one rudimentary; right molars with 5-6 spines, left
with 6-7 plus plume on both; palp article 2 with
small proximal and elongate apical inner setae,
basofacial formula on article 3 = 2 + 1 or 3 + 1.
Coxa 4 not broadened but smaller in relation to
coxa 1 than in female. long posterior setae on arti-
cle 2 of gnathopods 1-2 and pereopods 1-2 = 4-5-
3-4. Article 2 of only pereopods 4-5 narrower than
in female, anterior and apicolatcral spines on arti-
cle 3 of pereopod 5 reduced in length, article 4 of
pereopod 3 more produced distovemrally than in
female, article 6 narrowed, articles 5-6 of pereopod
4 more elongate than in female. Coxa 7 narrower
anteriorly and less sharply protuberant distroven-
trally than in female. Setal formulas, epimeron 1
anteroventral 6-7, postcroveniral = 2 (omitted in
illustration), epimeron 2 facial 6-7, epimeron 3
posterior one long. 2 short, facial 1, ventral =
4. Articulation of urosomitcs 1-2 complete. Spine
formulas of wropods, uropod I peduncle apico-
lateral 4-5, basofacial 5-7. medial 5-7 plus
apical enlarged, uropod '1 peduncle dorsal = 13-18,
dorsal spines on outer ramus of uropod 1 - 6-7, of
uropod 2 = 4-5, inner ramus of uropod I = 2-3, of
uropod 2 = 1-2 vestigial. Ventral spines on pedun-
cle of uropod 3 = 7-9, spine formula on article 1
of outer ramus = 1-1-1-1-1-2, setal formula =
1-1-1-1-1-1. Telson elongate, distal spines short-
ened.
ILLUSTRATIONS.?Upper and lower lips, maxillae,
maxilliped, and pereopods of the male resembling
those of the female and, except for minor differ-
ences mentioned in the text, are not illustrated;
male gnathopods are figured; normal female gna-
thopods and the atypical first gnathopod of the
holotype are drawn in outline only, that of the
holotype a medial view of the right member; draw-
ings of the urosome of both male and female speci-
mens show uropod 1 tilted slightly to reveal medial
spinules not seen in normal lateral view; mandibu-
lar protrusion at base of palp is as shown for B.
kokorus (25).
HOLOTYPE.?NMV, female "w," 6.7 mm.
TYPE-LOCALITY.?PPBES 223, 10 Mar 1971, Port
Phillip Bay, Victoria, Australia, 4 m, sand.
VOUCHER MATERIAL.?PPBES 224: male "u," 6.1
mm (illus); female "y," 7.9 mm (illus); male "a,"
5.6 mm (illus.). PPBES 230: female "n," 6.6 mm
(illus.). PPBES 231: female "f," 6.95 mm. PPBES
225: female "g," 6.95 mm.
RELATIONSHIP.?This species is very close to B.
NUMBER 245 SOS
FIGURE 146.?Birubius maamus, new species, holotype, female "w," 6.7 mm (a = male "a," 5.6
mm; n ?- female "n," 6.6 mm; v ? male "u," 6.1 mm; y ? female "y," 7.9 mm).
304 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
gambodeni (20) but can be distinguished by the
presence of one posteroproximal spine on article
5 of pereopods 1-2, by the shorter flagella and more
compact articles on antenna 1 and by the more
proximal placement of the ventral cycle of setae on
article 2 of antenna 1. The basofacial setal formula
on article 3 of the mandibular palp is very low com-
pared with B. gambodeni.
MATERIAL.?PPBES, 43 samples from 27 stations
(182); WPBES, one sample (1); CPBS, one sample
(1 doubtful).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 4-13 m, sand.
22. Birubius lowannus, new species
FIGURES 147-149
DESCRIPTION OF FEMALE.?Head about 18 percent
of total body length, greatest width about 67 per-
cent of length; rostrum constricted, narrow, reach-
ing middle of article 2 on antenna 1. Eyes medium,
clear of pigment. Article 1 on peduncle of antenna
1 about 1.5 times as long as wide, about 2.3 times
as wide as article 2, ventral margin with 4-5 setules,
weakly produced dorsal apex with 2 setules; article
2 about 0.8 times as long as article 1, with ventral
cycle of 6 setae; primary flagellum with 12-14 arti-
cles, about as long as peduncle, bearing aesthetascs;
accessory flagellum with 10-13 articles. Spine form-
ula on article 4 of antenna 2 = 1-3 4-4 or 1-3-5-4,
dorsal margin with notch bearing one seta and one
spine, ventral margin with 4 groups of 2 long to
medium setae, one ventrodistal medium spine; arti-
cle 5 about 0.7 times as long as article 4, facial
spine formula = 2, dorsal margin naked, ventral
margin with 2 sets of one seta, one setule, 3 ventro-
distal short to medium spines; flagellum about 1.5-
1.7 times as long as articles 4-5 of peduncle com-
bined, with 11-15 articles. Mandibles with me-
dium palpar hump; right incisor with 3 teeth; left
incisor with 3 teeth in 2 branches; right lacinia
mobilis bifid, distal branch simple, shorter than
proximal, tightly appressed, proximal branch sim-
ple, pointed, with marginal denticles; left lacinia
mobilis with 4 teeth and accessories; right rakers
7-8 plus 2-3 rudimentaries; left rakers 8-9 plus
3-4 rudimentaries; molar in form of elon-
gate plaque, right molar with 6-8 primarily long
spines plus one tiny disjunct, left molar with 8
primarily long spines plus one tiny spine disjunct,
each molar with plume; palp article 1 slightly elon-
gate, article 2 with one short inner apical seta and
2-3 other shorter inner setae, article 3 about 0.75
times as long as article 2, oblique apex with 6 spine-
setae, basofacial formula 0. Inner plate of max-
illa 1 somewhat thin but large, bearing one long
apical pluseta, one thinner apicomedial seta, 2 api-
colateral slightly shorter setae; palp article 2 with
6 apicalmedial marginal spines and 6-8 submar-
ginal setae. Plates of maxilla 2 extending sub-
equally, outer slightly narrower than inner, outer
with 3-4 apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with one large thick
apical spine, 3 apitofacial setae, 3-4 medial setae;
outer plate with 4 medial and apical spines, one
apicolateral seta; palp articles 1-2 with apicolateral
seta, medial margin of article 2 moderately setose,
article 3 with 3 facial setae, no lateral setae, nail of
article 4 medium, with one accessory setule. Coxa 1
strongly expanded distally, anterior margin weakly
concave; main ventral setae of coxae 1-4 = 6-6-5-0,
posteriormost seta of coxae 1-3 shortened; anterior
and posterior margins of coxa 4 weakly divergent,
posterior oblique, almost straight, posterodorsal
corner rounded, posterodorsal margin short,
straight, width-length ratio of coxa 4 = 13:15.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = l-4-4-(5-6), short posteriors =
2-3-3-(2^), long anteriors 3-(3-5)-O-O, short ante-
riors = 4-3-7-7. Gnathopods ordinary; width ratios
of articles 5-6 on gnathopods 1-2 = 22:33 and
24:36, length ratios = 72:62 and 64:60; palmar
humps ordinary, palms oblique; article 5 of gnatho-
pods 1-2 elongate, ovate, posterior margin rounded-
flat. Pereopods 1-2 similar; facial setae formula on
article 4 = 2 and 2, on article 5 = 3 and 3; main
spine of article 5 extending to M. 80 on article 6;
article 5 with one proximoposterior spine; spine
formula of article 6 = 4 + 4 plus middistal seta, 2
spines elongate; acclivity on inner margin of dactyls
of pereopods 1-2 weak, emergent setule long, mid-
facial pluseta ordinary. Coxae 5-7 posteroventral
seta formula = 4-10-9. Articles 4-5 of pereopods
3^4 broad, facial spine rows dense, facial ridge
formula on article 2 of pereopods 3-5 = 0-2-2, an-
terior ridge on pereopod 5 slightly shortened; width
ratios of articles 2, 4, 5, 6 of pereopod 3 = 44:49:
43:18, of pereopod 4 = 67:52=38:17, of pereo-
pod 5 = 88:23:20:10, length ratios of pereopod
NUMBER 245 305
S
FIGURE 147.?Birubius lowannus, new species, holotype, female "a," 5.80 mm (6 = male "b," 5.10
mm; c ? female "c," 4.50 mm; k = female "k," 6^5 mm; Y ? left lacinia mobilis).
306 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
3 = 75:42:44:44, of pereopod 4 = 86:57:39:51,
of pereopod 5 = 115:29:25:34; article 2 of pereo-
pod 5 reaching or exceeding middle of article 4;
article 2 of pereopod 3 slightly tapered distally;
medial apex of article 6 finely combed, bearing 2
small digital processes. Posteroventral corner of
epimeron 1 rounded, posterior margin weakly con-
vex, corner with setule (missing in illustration),
anteroventral margin with 5-6 short to medium
setae, posteroventral face with 2 long to short setae
set vertically, ventral seta short (unusual); postero-
ventral corner of epimeron 2 rounded, weakly pro-
tuberant, posterior margin straight, facial setae =
3-4, posteriormost pair set vertically; posteroven-
tral corner of epimeron 3 rounded, weakly and
doubly protuberant, with setule sinuses, posterior
margin straight, with 2 setule notches, ventral mar-
gin with one seta in middle, face with horizontal
row of 8 setae, epimeron 3 with setule on postero-
dorsal margin set in weak notch. Urosomite 1 with
ventral setule at base of uropod 1, articulation al-
most complete; urosomite 3 weakly protuberant
dorsally. Rami of uropods 1-2 with articulate en-
larged apical nails, outer ramus of uropod 1 with
4-6 dorsal spines, inner with 2, outer ramus of
uropod 2 with 3 dorsal spines, inner with one
dorsomedial spine; peduncle of uropod 1 with 2-3
apicolateral spines, no basofacial setae, medially
with one marginal seta and spinule, apically with
pair of ordinary spines; peduncle of uropod 2 with
7-9 dorsal spines, medially with one apical setule.
Peduncle of uropod 3 with 8-9 ventral spines, dor-
sally with one lateral spine, one medial spine and
setule; rami submasculine, inner extending to M.
80 on article 1 of outer ramus, apex with 3 setae,
medial margin with 2 setae, article 2 of outer
ramus ordinary, 0.24, bearing 2 medium setae, me-
dial margin of article 1 with one seta, lateral mar-
gin with 4 acclivities, spine formula = 2-2-2-2-2,
setal formula = 1-1-1-1-1. Telson ordinary, length-
width ratio =1:1, not fully cleft, each apex wide,
rounded, lateral acclivity deep, broad, bearing ordi-
nary lateral setule, spine next medial of length
equal to setule, midlateral setules diverse. Cuticle
with ordinary but sparse bulbar setules.
OBSERVATIONS (female).?Ommatidia larger than
in B. kokorus (25); articles 2-3 of antenna 2 with
sparse setular fringe dorsomedially, reminiscent of
male phoxocephalids; mandibular palp article 3
unusually short for members of B. gambodeni
group and coxae 1-4 with unusually high density
of facial setules.
Female "d,"' 7.3 mm, superfemale: Brood lamel-
lae normal; externally conforming to ordinary fe-
males in head, gnathopods, pereopods, epimera,
urosome, and telson but bearing only 9 spines on
each outer plate of maxilla 1; bearing 4 setae (in-
stead of one) on apex of palp article 2 on mandi-
ble; primary and accessory flagella of antenna 1
with 14 and 13 articles respectively; antenna 2 with
15 flagellar articles; peduncle of uropod 1 with 3
apicolateral spines, of uropod 2 with 9 dorsal
spines; each molar with very liny disjunct spine.
DESCRIPTION OF MALE.?Rostrum more sharply
constricted than in female. Article 2 of antenna 1
with 3 short ventral setae; primary flagellum with
11-12 articles, one calceolus each on articles 1-6
or 1-7, aesthetascs strongly developed. Facial spine
formula on article 4 of antenna 2 = 1-4?4-4, on
article 5 = 2, article 5 with 3 sets of dorsal male
setae and one calceolus, ventrodistal apex with 2
thin spines and one setule; flagellar formula =
(25-26), 2, 3. 5, 7 . . . 24 or 4, 5, 7, 9 . . . 23. Apical
seta on article 2 of mandibular palp longer than
in female, basofacial setal formula on article 3 =
1-2 (right only, left lost). Spine 2 on outer plate
of maxilla 1 unthickened. Coxa 4 broadened but
smaller in relation to coxa 1 than in female; ven-
tral setal formula of coxae 1-4 = 5-5 4 0. Article
2 of only pereopod 5 narrower than in female,
articles 5-6 of pereopod 4 and articles 3-5 of pe-
reopod 5 narrowed. Epimera 1-2 broadened, pos-
terior margin of epimeron 3 shortened; setal for-
mulas, epimeron 1 anteroventral = 3, posteroven-
tral = 2, epimeron 2 facial = 4, epimeron 3 poste-
rior = 3 setules, facial = 4. Spine formulas of uro-
pods, uropod 1 peduncle apicolateral = 2, basofa-
cial = 0, uropod 2 peduncle dorsal = 8, dorsal
spines on outer ramus of uropod 1 = 4, of uropod
2 = 3, inner ramus of uropod 1 = 2, of uropod
2 = 1 . Ventral spines on peduncle of uropod 3 = 5,
spine formula on article 1 of outer ramus = 1-1-
2-2-2-1, setal formula = 1-1-1-1-1-1. Telson
slightly elongate, distal spines shortened.
ILLUSTRATIONS.?Following parts not figured, re-
sembling those illustrated for species mentioned:
antenna 1 similar in proportions to B. kokorus (25)
and B. maamus (21); conformation of mandible as
in B. jirrandus (30); lower lip similar to that of
B. myallus (4); outer plate of female maxilla 1
NUMBER 245 307
FIGURE 148.?Birubius lowannus, new species, holotype, female "a," 5.80 mm (b ? male "b,"
5.10 mm; c - female "c," 4.50 mm; x - normal).
308 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 149.?Birubius lowannus, new species, holotype, female "a," 5.80 mm (b ? male "b,1
5.10 mm).
with one enlarged lateral spine as in B. myallus,
inner plate of female "c" (like "d") oriented to show
4 setae; inner plate of maxilliped displaced medi-
ally to show full length of outer plate; articles 3?7
of right female pereopod 5 illustrated to show nor-
mal condition in addition to complete left pereo-
pod with abnormally short articles; pereopod 4
of male "b" represented in outline only for com-
parison with female appendage.
HOLOTYPE.?NMV, female "a," 5.80 mm.
TYPE-LOCALITY.?CPBS 31E/1,25 Feb 1965, West-
ern Port, Victoria, Australia, 12 m, medium coarse
sand, a little fine mud.
VOUCHER MATERIAL.?CPBS 25N/5, male "b,"
5.10 mm (illus.); CPBS 25N/3, female "c," 4.50 mm
(illus.); PPBES 980/1, female "d," 7.3 mm; PPBES
NUMBER 245 309
980/2, female "k," 6.55 mm (illus.).
RELATIONSHIP.?Birnbius lowannus and B. kyee-
mus (23) form a subgroup within the greater B.
gambodeni group characterized by the following
combination of characters: organization of facial
setae on epimeron 3 into a horizontal row, absence
of long posterior setae and presence of one ventral
seta on epimeron 3, anteriorly crowded setae on
epimeron 2, shortened anterior facial ridge on arti-
cle 2 of pereopod 5, elongate wrists of the gnatho-
pods, and ordinary length of article 2 on the outer
rarnus of uropod 3.
The B. lowannus group differs from the strict
B. gambodeni subgroup in the elongate wrists of
the gnathopods, the ordinary article 2 of the outer
rannis on urojxxl 3 and in the organized setation
of epimeron 3. From the B. batei group (24-26)
the B. lowannus group (lifters in the absence of
long posterior setae and the strong facial row of
setae on epimeron 3 and the ordinary article 2 of
uropod 3. From the B. munggai group (27) the B.
lowannus group differs in the setal organization
on epimeron 3, the short anterior ridge on article
2 of pereopod 5, and the ordinary article 2 of uro-
pod 3.
MATERIAL.?CPBS, 12 samples from 6 stations
(14); WPBES, 9 samples from 6 stations (20);
PPBES, 8 samples from 2 stations (16).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 7-31 m, sand.
23. Birubius kyeemus, new species
FIGURES 150, 151
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about 62
percent of length; rostrum constricted, narrow,
elongate, reaching middle of article 2 on antenna
1. Eyes small, largely occluded with pigment. Arti-
cle 1 on peduncle of antenna 1 about 1.5 times as
long as wide, about twice as wide as article 2, ven-
tral margin with about 6 setules, weakly produced
dorsal apex with 2 setules; article 2 about 0.60
times as long as article 1, with ventral cycle of 5
setae; primary flagellum with 10-11 articles, about
0.8 times as long as peduncle, bearing several short
aesthetascs; accessory flagellum with 8-9 articles.
Spine formula on article 4 of antenna 2 = 1-3-4?4,
dorsal margin with subproximal spine, ventral
margin with 4 groups of 2 long to medium setae,
one ventrodistal medium spine; article 5 about 0.75
times as long as article 4, facial spine formula =
1-3, dorsal margin naked, ventral margin with 2
sets of 3-4 long to medium setae, 2 ventrodistal
long to medium spines; flagellum about 1.7 times
as long as articles 4-5 of peduncle combined, with
12-14 articles. Mandibles with medium palpar
hump; right incisor with 3 teeth; left incisor with
3-4 humps in 2 branches, right lacinia mobilis
bifid, distal branch shorter than proximal, both
branches simple, pointed, proximal with marginal
denticles; left lacinia mobilis with 5 teeth plus one
accessory tooth, middle teeth shortened; right rak-
ers 6; left rakers 8; molar in form of elongate
plaque, each molar with 5 primarily long spines,
none disjunct, each molar with plume; palp article
1 slightly elongate, article 2 with one long inner
apical seta and 2 other shorter inner setae, article
3 about 0.75 times as long as article 2, oblique
apex with 8 spine-setae, basofacial formula = 0.
Inner plate of maxilla 1 elongate, thin, bearing one
long apical pluseta, one shorter similar apicomedial
seta, 2 apicolateral somewhat enlarged setae; palp
article 2 with one apical spine and 4 apicolateral-
apical-medial setae. Plates of maxilla 2 extending
equally, of equal breadth, outer with 3 apicolateral
setae, inner with one medial seta. Inner plate of
maxilliped with one large thick apical spine, 2
apicofacial setae, 4 medial setae, outer plate with
4?5 medial and apical spines, one apicolateral seta
and 2 setules; palp articles 1-2 with apicolateral
seta, medial margin of article 2 moderately to
weakly setose, article 3 protuberant, with 3 facial
setae, no lateral setae, nail of article 4 medium,
with 2 accessory setules. Coxa 1 expanded distally,
anterior margin weakly concave; main ventral setae
of coxae 1^ 4 = (5-6)-(5-6)-(5-6)-0, posteriormost
seta of coxae 1-3 shortened; anterior and posterior
margins of coxa 4 divergent, posterior margin
oblique, almost straight, posterodorsal corner
rounded, posterodorsal margin ordinary, straight,
width-length ratio of coxa 4 = 10:11. Long pos-
terior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 1-3-2-3, short posteriors = 0-0-3?
(2-3), long anteriors = 0-4-0-0, short anteriors =
2-3-3-4. Gnathopods ordinary; width ratios of arti-
cles 5-6 on gnathopods 1-2 = 22:32 and 24:34,
length ratios = 66:59 and 59:58; palmar humps
ordinary, palms weakly oblique; article 5 of gnath-
310
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s
FIGURE 150.?Birubius kyeemus, new species, holotype, female "a," 5.94 mm.
opods 1-2 elongate, ovate, posterior margin flat.
Pereopods 1-2 similar, facial setae formula on arti-
cle 4 = 3 and 2, on article 5 = 3 and 4; main spine
of article 5 extending to M. 75 on article 6, article
5 with one proximoposterior spine; spine formula
on article 6 = 4 + 5, some elongate, plus tiny mid-
distal seta (omitted from illustration); acclivity on
inner margin of dactyls of pereopods 1-2 sharp,
produced as tooth, emergent setule short, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule
formula = 4-5-5. Articles 4-5 of pereopods 3-4
broad, facial spine rows dense, facial ridge formula
on article 2 of pereopods 3-5 = 0-2-2, anterior on
pereopod 5 short; width ratios of articles 2, 4, 5, 6
NUMBER 245 SH
FIGURE 151.?Birubius kyeemus, new species, holotype, female "a," 5.94 mm.
312 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
of pereopod 3 = 42:47:41:18, of pereopod 4 =
71:51:33:17, of pereopod 5 = 74:22:19:10, length
ratios of pereopod 3 = 74:26:39:42, of pereopod
4 = 88:51:42:59, of pereopod 5 = 100:28:24:34;
article 2 of pereopod 5 reaching middle of
article 5, ventral margin with one medium
seta plus setules; medial apex of article 6
scarcely combed, straight. Posteroventral corner of
epimeron 1 rounded, weakly protuberant, posterior
margin weakly convex, with setule notch, antero-
ventral margin with 3-4 medium setae, posteroven-
tral face with 2 long setae set vertically; postero-
ventral corner of epimeron 2 rounded, posterior
margin weakly convex, with setule, facial setae =
4-5 crowded anteriorly, posterior pair set verti-
cally; posteroventral corner of epimeron 3 rounded,
weakly protuberant, with setule sinus above and
below, posterior margin almost straight, naked,
ventral margin naked, midface with scarcely
oblique row of 3-4 setae plus one seta near ventral
margin; epimera 1-2 with weak setule notch pos-
terodorsally. Urosomite 1 with ventral setule at
base of uropod 1, articulation line almost absent;
urosomite 3 scarcely protuberant dorsally. Rami of
uropods 1-2 with articulate enlarged apical nails,
outer ram us of uropod 1 with 2-3 dorsaJ spines,
inner with 2, outer ramus of uropod 2 with 2 dor-
sal spines, inner with 2 dorsomedial spines; pe-
duncle of uropod 1 with 2 apicolateral spines and
2 basofacial setae, medially with one marginal seta
and apical enlarged spine; peduncle of uropod 2
with 5 dorsal spines, medially with one setule. Pe-
duncle of uropod 3 with 5 ventral spines, dorsally
with one lateral spine, one medial setule; rami
feminine, inner extending to M. 57 on article 1
of outer ramus, apex with 3 setae, medial and lat-
eral margins naked, article 2 of outer ramus ordi-
nary to elongate, 0.25, bearing 2 long setae, sub-
apicomedial margin of article 1 with one seta, lat-
eral margin with 3-4 acclivities, spine formula =
1-2-2-2-2 or 1-2-2-2, setal formula = 0. Telson
short, length-width ratio = 51:57, not fully cleft,
each apex wide, rounded, lateral acclivity broad,
shallow, bearing ordinary lateral setule, spine next
medial shorter than setule, midlateral setules di-
verse. Cuticle with ordinary sparse bulbar setules.
HOLOTYPE.?AM, female "a," 5.94 mm.
TYPE-LOCALITY.?EBS 148, 12 Jan 1973, 500 yards
off Towra Point, Botany Bay, New South Wales,
Australia, 3 m, sand and shell grit.
VOUCHER MATERIAL.?Type-locality: female "b,"
5.42 mm; female "c," 5.36 mm. Male unknown.
RELATIONSHIP.?This species belongs in a group
with Birubins lowannus (22) and shares with that
species all the character differences from other spe-
cies groups discussed in the "Relationship" section
of that species.
Birubius kyeemus differs from B. lowannus in
the poor articulation line of urosomite I. the pres-
ence of basofacial setae on uropod I, shorter spines
on the telson, the simultaneous presence of more
setae on epimeron 2 and fewer on epimeron 3,
the presence of 2 (not 1) dorsal spines on the inner
ramus of uropod 2, a much shorter inner ramus of
fcmala uropod 3 and the disjunct pairing of 2
facial setae set proximally on palp article 3 of the
maxilliped.
MATERIAL.?EBS, one sample (3).
DISTRIBUTION.?New South Wales, Botany Bay,
Towra Point, 3 in, sand and shell grit.
24. Birubius bateif (Haswell), new combination
FIGURES 152-155
Phoxus Batei Haswell, 1879:259-260, pi. 9: fig. S; 1882:237-
238.
Pontharpinia rostrata.?Stebbing, 1906:146-147 [in part]:
1910:635.
NOMENCLATURE.?The so-called type material of
this species (AM, P.3438, Port Jackson) is con-
founded. The specimens in the collection are
Matong matong, new species, and clearly not what
Haswell described and illustrated (Barnard and
Drummond, 1976, erroneously labeled this taxon
as "Birubius sp."). There is strong evidence that
the material under consideration was labeled as
"Type" by a person other than Haswell more than
20 years after his publication (see J. L. Barnard,
1974). We therefore reject that material as being
the legitimate type of Birubius batei. On the other
hand we hesitate in creating a neotype for this
species; instead, we have designated a "Main
Voucher." We have material, from Botany Bay, a
locality near Port Jackson, that fits Haswell's
meager description, but believe that a precise sur-
vey of the Port Jackson area should be undertaken
to find what lives there today. Another of our spe-
cies, B. jirrandus (30), also fits the description of
B. batei but has not, as yet, been found in the
NUMBER 245 313
S
FIGURE 152.?Birubius bateii (Haswell), new combination, main voucher, female "f," 5.39 mm
(z ? male "z," 550 mm; Y = left lacinia mobilis).
314 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
vicinity of Port Jackson, New South Wales.
DESCRIPTION OF FEMALE.?Head about 25 per-
cent of total body length, greatest width about 65
percent of length; rostrum weakly constricted, nar-
row, reaching middle of article 2 on antenna 1.
Eyes medium, largely occluded with pigment. Arti-
cle 1 on peduncle of antenna 1 about 1.5 times as
long as wide, about 2.4 times as wide as article 2,
ventral margin with about 10 setules, weakly pro-
duced dorsal apex with 3-4 setules (only 2 showing
in illustration); article 2 about 0.55 times as long
as article 1, with ventral proximal cycle of 7 setae;
primary flagellum with 12 articles, about 0.85 times
as long as peduncle, lacking aesthetascs; accessory
flagellum with 10 articles. Spine formula on article
4 of antenna 2 = 1-3-4-3, dorsal margin with notch
bearing 3 setae and spines, ventral margin with C
groups of 2 long to short setae, one ventrodistal
long spine; article 5 about 0.8 times as long as
article 4, facial spine formula = 1-2-2, dorsal mar-
gin bearing group of setules, ventral margin with
4 sets of 2-3 long to short setae, 3 ventrodistal long
to medium spines; flagellum about 1.3 times as
long as articles 4-5 of peduncle combined, with 13
articles. Mandibles with medium palpar hump;
right incisor with 3 teeth and notch; left incisor
with 3 teeth in 2 branches; right lacinia mobilis
bifid, distal branch simple, shorter than proximal,
proximal branch simple, pointed; left lacinia mo-
bilis with 4 teeth plus one accessory tooth; right
rakers 8; left rakers 9; molar in form of elongate
plaque, molars with 8 primarily long spines, no
spine disjunct, each molar with plume; palp article
1 short, article 2 with one long inner apical seta
and 3 other short inner setae and 2 outer setae,
article 3 about as long as article 2, oblique apex
with 10 spine-setae, basofacial formula = 0. Inner
plate of maxilla 1 large, bearing one long apico-
facial pluseta, one shorter similar apicomedial seta,
2 apicolateral much shorter setae; palp article 2
with 2-3 apicalmedial marginal spines and 5-6
submarginal setae. Plates of maxilla 2 extending
equally, of equal breadth, outer with 6 apicolateral
setae, inner with 3 medial setae. Inner plate of
maxilliped with one large thick apical spine, 2
apicofacial setae, 4 medial setae; outer plate with
6 medial and apical spines, 2 apicolateral setae;
palp article 1 with 2 apicolateral setae, article 2
with 3 groups of one apicolateral seta each, medial
margin of article 2 moderately setose, article 3 pro-
duced, with 4-5 facial setae. 2 lateral setae, nail of
article 4 medium, with 2 accessory setules. Coxa 1
scarcely expanded distally, anterior margin almost
straight, main ventral setae of coxae 1-4 = (5-6)-
7-7-0, posteriormost seta of coxae 1-2 slightly
shortened; anterior and posterior margins of coxa
4 divergent, posterior margin oblique, convex, pos-
terodorsal corner rounded, posterodorsal margin
short, almost straight, width-length ratio of coxa
4 = 7:8. Long posterior setae on article 2 of gnath-
opods 1-2 and pereopods 1-2 = 3-(5-6)-4-6, short
anteriors = (10-11 )-(8-9)-5-3. long anteriors = 3-
(13-14)-O-O, gnathopod 1 with 3 long basofacial
setae. Gnathopods ordinary; width ratios of articles
5-6 on gnathopods 1-2 - 24:30 and 26:33. length
ratios = 65:<>() and 58:57; palmar humps ordi-
nary, palms oblique; article 5 of gnathojxxls 1-2
elongate, ovate, posterior margin flat. Pcreopods
1-2 similar; pereopod 1 with supernumerary apico-
facial setae on article 5, other facial setae formula
on article 4 = 4 and 3, on article 5 = 5 and (i; main
spine of article 5 extending to M. 75 on article 6,
article 5 with 2 proximoposterior spines; spine for-
mula of article 6 = 5 + 6 plus middistal seta; ac-
clivity on inner margin of dactyls of pereopods 1-2
weak, emergent setule long, midfacial pluseta ordi-
nary. Coxae 5-7 posteroventral setule formula =
4?3-4. Articles 4-5 of pereopods 3?4 broad, facial
spine rows moderately developed, facial ridge for-
mula on article 2 of pereopods 3-5 = 0-2-2, ante-
rior ridge of pereopod 5 long, posterior ridge abut-
ting; width ratios of articles 2, 4, 5, 6 of pereo-
pod 3 = 41:40:34:16, of pereopod 4 = 67:39:26:14,
of pereopod 5 = 77:26:19:9, length ratios of pe-
reopod 3 = 78:34:43:42, of pereopod 4 = 90:52:41:
54, of pereopod 5 = 102:36:28:20; article 2
of pereopod 5 reaching apex of article 4 (dis-
counting special lobe); article 4 of pereopods 3 and
5 with especially elongate posterodistal lobe; me-
dial apex of article 6 truncate, uncombed. Postero-
ventral corner of epimeron 1 rounded, posterior
margin convex, bearing setule, corner with setules,
anteroventral margin with 6 short setae, postero-
ventral margin with 2 long setae; posteroventral
corner of epimeron 2 rounded, weakly protuberant,
posterior margin undulant, with setule, facial
setae = 8-10, posteriormost pair set vertically; pos-
teroventral corner of epimeron 3 with small tooth,
posterior margin straight, serrate, 7-8 setose, upper-
most seta tiny, ventral margin with 3-4 posterior
NUMBER 245 315
s
FIGURE 153.?Birubius batci? (Haswell), new combination, main voucher, female "f," 5.39 mm
(z ? male '%" 5.50 mm).
316 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
setae. Urosomite 1 with 2 long ventral spines at
base of uropod 1, articulation line broken in
middle; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1-2 with articulate enlarged
apical nails, rami of uropods 1-2 lacking accessory
nails, outer ramus of uropod 1 with 6-10 dorsal
spines, inner with 2-4, outer ramus of uropod 2
with 4-8 dorsal spines, inner with 1-2 dorsomedial
spines; peduncle of uropod 1 with 4-5 apicolateral
spines and 4-5 basofacial setae, medially with 6 mar-
ginal setae and spines becoming thicker apically;
peduncle of uropod 2 with 10-14 dorsal spines,
medially with one small apical spinule. Peduncle of
uropod 3 with 6 ventral spines, dorsally with one
lateral spine, one medial spine; rami feminine, inner
extending to M. 75 on article 1 of outer ramus, apex
with 3 setae, medial margin with 3 setae, article 2 of
outer ramus ordinary, 0.17, bearing 2 long setae,
medial margin of article 1 with 2 setae, lateral mar-
gin with 5 acclivities, spine formula = 1-2-2-2-
2-2, setal formula = 0-1-1-1-1-1. Telson ordinary,
length-width ratio = 14:13, not fully cleft, each
apex of medium width, rounded, lateral acclivity
shallow, bearing ordinary lateral setule, spine next
medial of length equal to setule, midlateral setules
diverse. Cuticle with ordinary sparse bulbar setules,
surface bearing fine striations in form of linear
fingerprint pattern.
DESCRIPTION OF MALE.?Rostrum shorter than in
female. Antenna 1 with one calceolus each on arti-
cles 1-4. Facial spine formula on article 4 of an-
tenna 2 = 1-2-4-3, on article 5 = 2; article 5 with
one dorsal set of male setae and one calceolus, ven-
trodistal apex with 2 thin spines and one setule,
flagellar formula = 27, 2, 4, 6 . . . 24. Right and
left rakers 8; basofacial setal formula of article 3 on
mandibular palp = 1-2. Palp of maxilla 1 with 2
spines and 4 setae. Outer plate of maxilliped with
one apicolateral seta. Coxa 4 like that of female
but smaller in relation to coxa 1 than in female;
ventral setal formula of coxae 1-4 = 4-6-6-0, setae
short. Long posterior setae on article 2 of gnatho-
pods 1-2 and pereopods 1-2 = 2-(2-3)-2-3 plus 2
basofacial setae on gnathopod 1, long anteriors =
3-7-0-0. Facial and setal-spine formulas of pereo-
pods 1-2 on article 4 = 3 and 2, on article 5 = 5
and 5, on article 6 = 4 and 5 plus middistal seta.
Article 2 of only pereopods 3 and 5 narrower than
in female, pereopod 5 larger in relation to pereo-
pod 3 than in female. Epimera 1-3 broadened, pos-
terior margin of epimeron 3 shortened; setal form-
ulas, epimeron 1 anteiovemral = 5, posteroven-
tral = 2, epimeron 2 facial = 7, epimeron 3 poste-
rior = 8, facial = 1-2. Articulation lines of urosome
strong, ventrolateral spine of urosomite 1 very
small, dorsal base of urosomite 3 with hump. Spine
formulas of uropods, uropod 1 peduncle apico-
lateral = 4-5, basofacial = 6, medial = 4 plus
apical enlarged, uropod 2 peduncle dorsal = 11-14,
dorsal spines on outer ramus of uropod 1 = 6-7, of
uropod 2 = 5-6, inner ramus of uropod 1 = 2-3, of
uropod 2 = 2 shortened, fully medial. Ventral
spines on peduncle of uropod 3 = 6, spine formula
on article 1 of outer ramus = 1-1-2-2-2-2, setal
formula = 1-1-1-1-1-1. Telson slightly elongate,
distal spines shortened.
OBSERVATIONS.?Dwarf males from Western Aus-
tralia, VVAM 414-73 (index "273'): Male "a." 3.31
mm, and male "b," 3.50 mm: nail on dactyl of
maxilliped elongate, 60 |>ercent as long as article 4;
article 6 of gnathopod 2 stouter than in typical
phenotype. Other descriptive features of male "a":
primary flagellum of antenna 1 with 10 articles, ac-
cessory flagellum with 7, primary with one calceolus
each on articles 1-4. One aesthetasc each on articles
1-8; antenna 2 flagellar formula = 23, 2, 4, 6 . . .
20; article 4 of peduncle with spine formula of
3-4-3, article 5 with 2 facial spines and 2 sets of
posterior setal spines; right lacinia mobilis of
normal bifid form, mandibular palp article 2 with
3 medial setae and 2 apicomedial setae, article 3
with 7 apical setae, basofacial formula = 2-1; palp
of maxilla 1 with 4 spines and 3 setae; setal form-
ula of coxae 1^ 1 = 4-5-5-0; long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
1-1-2-2, long anteriors = (2-3)-4-O-O, short ante-
riors = (O-l)-3-3-3, no others, facial setae on arti-
cles 4 and 5 of pereopods 1-2 = 2 and 3, spine
formula of article 6 = 3 + 4 plus middistal seta;
anterior ridge on article 2 of pereopod 5 of short
form seen in Birubius jirrandus (30), pereopod 5
with 6 posterior teeth on article 2; epimeron 1 with
4 anteroventral setae, 2 posteroventral setae, epi-
meron 2 with 4 horizontal setae and pair set ver-
tically, epimeron 3 with 6 posterior setae and one
or no posteroventral seta; peduncle of uropod 1
with 2 basofacial setae, with 4 apicolateral spines,
outer ramus with 6 dorsal spines, inner ramus with
2, peduncle of uropod 2 with 9 dorsal spines, outer
ramus with 3, inner with one; outer ramus of uro-
NUMBER 245 517
R3
FIGURE 154.?Birubius bateif (Haswell), new combination, main voucher, female "f," 5.39 mm
(z ? male "z," 530 mm).
pod 3 with 4 acclivities, spine formula = 1-1-1-1-2,
setal formula = 1-1-1-1-1. Male "b": epimeron 1
with 4 anteroventral and 3 posteroventral setae,
epimeron 2 with 5 horizontal setae and pair set
vertically, epimeron 3 with 7 posterior and 2 pos-
teroventral setae; peduncle of uropod 1 with 5 baso-
facial setae, peduncle of uropod 2 with 10 dorsal
spines, outer ramus with 4; other counts similar to
male "a".
Our southeastern Australian material of Birubius
bateif is mostly represented by giant specimens,
whereas the dwarf western Australian males may
represent a more average size for this species. Fea-
tures of subspecific value between the two sets of
specimens collected almost 2,000 miles apart in-
clude spine counts, the short ridge on pereopod 5,
and the broadened hands on the gnathopods.
Males of WAM 414-73 (index "274"):Male "m,"
4.80 mm: subbifid apical branch of right lacinia
mobilis (see illustration); antenna 2 normal; nail
on maxillipedal palp almost 40 percent as long as
article 4, thus slightly elongate; hand of gnathopod
318 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
1 almost normal, that of gnathopod 2 stout (see
illustration); pereopod 5 normal; epimeron 1 with
5 anteroventral setae, 2 posteroventral submarginal
setae, epimeron 2 with 5 horizontal and one dor-
sally thrust setae, epimeron 3 with 6 posterior and
no ventral setae; peduncle of uropod 1 with 4-5
basofacial setae, 5 apicolateral spines, outer ramus
with 6 spines, inner with 2, peduncle of uropod 2
with 11 spines, outer ramus with 3 dorsal spines
and one spine fully inserted (non-emergent) in
body of ramus, inner ramus with 2 spines; lateral
apex of peduncle on uropod 3 with 2 spines, outer
ramus with spine formula of 1-1-2-2-2-2, setal
formula of 1-1-1-1-1-1; abnormalities are maxil-
lipedal nail, gnathopod 2, outer ramus of uropod 2,
peduncle of uropod 3. Male "n": right lacinia
mobilis normal; both hands of gnathopods stout;
outer ramus of uropod 2 with 5 normal spines, in-
ner with one; spine formula on outer ramus of
uropod 3 = 2-1-1-1-2, setae = 0-2-1-1-1; abnor-
malities are gnathopods and outer ramus of uropod
3. Male "p": Only hand of gnathopod 2 stout.
Males of WAM 414-73 (index "275"): Male "q":
right lacinia mobilis normal; maxillipedal nail
normal; hands of both gnathopods slightly stout-
ened; anterior ridge of pereopod 5 normal; epi-
meron 1 with 5 anteroventral setae, with 2 postero-
ventral setae immediately adjacent to anteroventral
set (abnormal), epimeron 2 with 6 horizontal setae
and one thrust dorsally, epimeron 3 with 4 poste-
rior setae confined ventrally, one posteroventral
seta, one ventral seta-spine; peduncle of uropod 1
with 6 basofacial setae, lateral apex with 6 spines
(one of these disjunct towards base), outer ramus
with 5 spines, inner with one (abnormal), peduncle
of uropod 2 with 11 spines, outer ramus with 4, in-
ner with one; spine formula on outer ramus of
uropod 3 = 1-1-1-2-2-2, setae = 0-1-1-1-1-1.
Other males: epimeron 1 normal.
Abnormalities in Western Australian Males: Re-
combinations or gene flow in the direction of Biru-
bius jirrandus (30) are seen in the shortened ante-
rior ridge of pereopod 5 in "273" males, in the
spine reduction on the inner ramus of uropod 1 on
"275" males; but other abnormalities in maxil-
lipedal nail, setal distribution on epimera 1-2, and
right lacinia mobilis of one "274" male, all of which
occur inconsistently, suggest that the two main
differences listed in the discussion of this species
and B. jirrandus are valid.
DESCRIPTION OF JUVENILE (juvenile "k," 2.26 mm).
?Head similar to that of adult but somewhat thin-
ner from dorsal view. Antenna 1 with 8 articles in
primary flagellum, 5 in accessory flagellum. Spine
formula on article 4 of antenna 2 = 1-3-3-3, dorsal
margin with 2 setae in notch, article 5 with 2
facial spines and 2 posterior sets of setae; flagellum
7-articulate. Upper lip and epistome lacking articu-
lation. Right lacinia mobilis bifid, both branches
conical; right mandible with 5 raker spines; molar
with 3 spines, none disjunct (left mandible not exam-
ined); palp article 3 lacking basofacial setae. Lower
lip and plates of maxilla 1 not examined; palp of
maxilla 1 with 2 spines and 3 setae. Inner plate of
maxilla 2 with one medial seta; outer plate with
one apicolateral seta. Inner plate of maxilliped
with one apical spine and one apicoventral seta, 2
medial and 2 dorsofacial setae; outer plate with 4
spines, no lateral setae; article 3 of palp with 2
facial setae. Setae of coxae 1-4 = 4-3-3-0. Article
2 of gnathopods 1-2 with 0 and 2 long posterior
(see illustrations of gnathopods). Setal and spine
formulas of pereopods 1-2, article 2, posterior long
seta = 1, article 4 facial = 4, article 5 facial = 2,
article 6 spines = 3 + 3 plus one seta, article 5 also
bearing one proximal spine. Pereopods 3-4 gen-
erally similar to those of adult but with fewer
spines and setae. Article 2 of pereopod 5 (illustrated)
with stronger ventral setae than in adult (generally
typical of juveniles in certain species). Epimeron 1
(illustrated) with one anteroventral seta, one pos-
teroventral seta; epimeron 2 with 4 facial setae in
tandem; epimeron 3 with 3 posterior setae and 2
setules, no facial setae. Urosomite 1 with one large
ventral spine at base of each uropod 1 (not in
adult). Uropod 1 with a basofacial seta, 1-2 apico-
lateral spines, outer ramus with 3 dorsal spines,
inner with one dorsal spine; uropod 2 with 4-5
peduncular spines, outer ramus with 2 dorsal
spines, inner with one dorsal spine. Uropod 3 and
telson as illustrated.
Identification of juvenile with adult is strong but
confounded by the relatively great length of the
ventral setules on article 2 of pereopod 5. This
anomaly can be shown in other species to be char-
acteristic of youthful specimens in contrast to
adults.
ILLUSTRATIONS.?The paired basofacial setae on
article 3 of the maxillipedal palp in female "f" is
an abnormal condition.
NUMBER 245 319
mGl nJ mG2
FIGURE 155.?Birubius bateif (Haswell), new combination, juvenile "k," 2.26 mm (m = male
"m," 4.80 mm; n ? male "n," 4.70 mm).
MAIN VOUCHER.?AM, female "f," ovigerous, 5.39 "d," 6.10 mm. EBS 196: male "e," 5.62 mm. WAM
mm. 414-73 (divided into 3 subsamples with indices):
LOCALITY.?EBS 200, 11 Apr 1973, off Towra "273," male "a," 3.31 mm, male "b," 3.50 mm;
Beach, Botany Bay, New South Wales, Australia, "274," male "m," 4.80 mm (illus.), male "n," 4.70
2.7 m, sand. mm (illus.), male "p," 4.20 mm; "275," male "q,"
VOUCHER MATERIAL.?EBS 362: male "z," 5.50 4.36 mm; plus 3 other males. JLB AUS 13: juvenile
mm (illus.); female "c," 6.28 mm. EBS 199: female "k," 2.26 mm (illus.).
320 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
RELATIONSHIP.?Because the combination of non-
crowded setae on epimeron 2, strong basofacial
setae on uropod 1 and elongate wrists on the gna-
thopods prevents an evolutionary flow from the
other 3 subgroups in this division, Birubius batei?
and B. jirrandus (30) form a pair distinct from
the other members of this division and apparently
derive not from any of the other subgroups but
directly from their hypothetical ancestor. Birubius
batei? and B. jirrandus also differ from the other
group in the absence of fully facial setae on epi-
meron 3 and in the presence of ventral setae,
although the latter are almost facial.
In other ways this species appears very close to
the B. kokorus group (25-27) but differs in the
more numerous basofacial setae on uropod 1 and /or
in the longer telsonic spines. From the B. lowannus
group (22-23), B. batei? differs in the longer ante-
rior ridge of pereopod 5, the uncrowded setae of
epimeron 2, and the shorter article 2 on the outer
ramus of uropod 3. From the B. gambodeni group
(20-21) B. batei? differs in the more elongate wrists
of the gnathopods and the longer anterior ridge on
pereopod 5.
MATERIAL.?EBS, 20 samples (52); WAM, one
sample (11); JLB AUS, one sample (1).
DISTRIBUTION.?New South Wales: Botany Bay,
Lake Macquarie, and Jervis Bay, 0.3-6.0 m, coarse
sand. Western Australia: Barrow Island, presum-
ably surface nekton; and Middleton Beach, near
Albany, intertidal.
25. Birubius kokorus, new species
FIGURES 156-158
DESCRIPTION OF FEMALE.?Head about 17 per-
cent of total body length, greatest width about 63
percent of length; rostrum constricted, narrow, of
medium length, reaching middle of article 2 on
antenna 1. Eyes medium, largely occluded with pig-
ment. Article 1 on peduncle of antenna 1 about
1.3 times as long as wide, about 2.2 times as wide
as article 2, ventral margin with few setules, un-
produced dorsal apex with one setule; article 2
about 0.75 times as long as article 1, with ventral
cycle of 4-7 setae; primary flagellum with 11-14
articles, about 0.75-0.87 times as long as peduncle,
bearing long aesthetascs distally; accessory flagellum
with 9-12 articles. Spine formula on article 4 of
antenna 2 = 1-3^1-5 or 1-3-5-5 or 1-3-6-5 or
1_3_5_4( dorsal margin with notch bearing 3 setae,
ventral margin with 4-6 groups of 3-4 long to me-
dium setae, one ventrodistal long spine; article 5
about 0.75 times as long as article 4, facial spine
formula = 1-3 or 1-4 or 1-2-2. dorsal margin
naked, ventral margin with 4 sets of 2 long and
short setae, 2 ventrodistal long to medium spines;
flagellum about 1.2 times as long as articles 4-5
of peduncle combined, with 12-14 articles. Man-
dibles with medium palpar hump; right incisor
with 3 teeth; left incisor with 3 teeth in 2 branches;
right lacinia mobilis bifid, distal branch simple,
shorter than proximal, tightly appressed, proximal
branch simple, pointed, with marginal denticles;
left lacinia mobilis with 4 teeth; right rakers 8-9
plus 1-2 rudimentaries; left rakers 8-10 plus 1-2
rudimentaries; molar in form of elongate plaque,
right and left molars with 7 primarily long spines,
no spine disjunct, each molar with plume; palp
article 1 short, article 2 with one medium inner
apical seta and 3-4 other shorter inner setae, arti-
cle 3 about as long as article 2, oblique apex with
10 spine-setae, basofacial formula = 0-1 or 0-2.
Inner plate of maxilla 1 large, bearing one long
apical pluseta, one shorter similar apicomedial seta,
2 apicolateral much shorter setae (one hidden in
illustration); palp article 2 with 3?4 apical medial
marginal spines and 4?5 submarginal setae. Plates
of maxilla 2 extending equally, plates equally
broad, outer with 5-6 apicolateral setae, inner with
one medial seta. Inner plate of maxilliped with
one large thick apical spine, 2 apicofacial setae, 4
medial setae, outer plate with 6-7 medial and api-
cal spines, 2 apicolateral setae; palp article 1 with
apicolateral seta, article 2 with 1-2 groups of 1-2
apicolateral setae, medial margin of article 2
strongly setose, article 3 protuberant, with 5 facial
setae, no lateral setae, nail of article 4 short, with
1-2 accessory setules. Coxa 1 expanded distally,
anterior margin weakly concave; main ventral setae
of coxae l^t = 10-9-9-0, posteriormost seta of coxae
1-2 slightly shortened; anterior and posterior mar-
gins of coxa 4 weakly divergent, posterior margin
convex, posterodorsal corner rounded, posterodorsal
margin short, concave, width-length ratio of coxa
4 = 14:15. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 3-(5-7)-5-6,
long anteriors = H?I?0-0, others not counted.
Gnathopods generally ordinary; width ratios of
NUMBER 245 321
nS
FIGURE 156.?Birubius kokorus, new species, holotype, female "a," 7.7 mm (n = female "n," 8.2
mm; q ? male "q," 6.3 mm; y ? female "y," 6.3 mm).
322 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
articles 5-6 on gnathopods 1-2 = 25:32 and 26:33,
length ratios = 54:61 and 49:60; palmar humps
ordinary, palms oblique; article 5 of gnathopod 1
ovate, posterior margin flat; article 5 of gnathopod
2 ovate, posterior margin rounded, lobate. Pereo-
pods 1-2 similar; facial setae formula on article
4 = 4?5, on article 5 = 6; main spine of article 5
extending to M. 90 on article 6, article 5 with 2
proximoposterior spines; spine formula of article
6 = 3 + 4 (rarely 4 + 5) plus middistal seta; ac-
clivity on inner margin of dactyls of pereopods 1-2
weak, produced as tooth, emergent setule long,
midfacial pluseta ordinary. Coxae 5-7 posteroven-
tral setule formula = 4-4-(3-4). Articles 4-5 of pe-
reopods 3-4 broad, facial spine rows moderately
developed, facial ridge formula on article 2 of
pereopods 3-5 = 0-2-2, anterior ridge on pereopod
5 slightly shortened; width ratios of articles 2, 4,
5, 6 of pereopod 3 = 45:42:36:17, of pereopod
4 = 71:42:32:14, of pereopod 5 = 87:27:23:10,
length ratios of pereopod 3 = 72:33:33:43, of
pereopod 4 = 88:53:43:52, of pereopod 5 =
115:29:26:30; article 2 of pereopod 5 reaching
middle of article 5; medial apex of article 6 finely
combed, bearing 4 obsolescent digital processes.
Posteroventral corner of epimeron 1 rounded, pro-
tuberant, posterior margin convex, corner with
setule, anteroventral margin with 8-10 medium
setae, posteroventral face with 1-2 long setae; pos-
teroventral corner of epimeron 2 rounded, weakly
protuberant, guarded by setule sinus, posterior
margin weakly convex, facial setae = 10-14, pos-
teriormost pair or triad of main group set verti-
cally, with additional disjunct seta; posteroventral
corner of epimeron 3 weakly protuberant, with
setule sinus, with small tooth, posterior margin
weakly sinuous, with several setule notches and 3-4
long setae, ventral margin with 2-3 setules mainly
posterior, face with 2 oblique horizontal rows of
2-3 setae or variants (see illustrations) near ventral
margin in middle; epimeron 3 with setule on
posterodorsal margin set in weak notch. Urosomite
1 with ventral setule at base of uropod 1, (not
showing in illustration), articulation line almost
complete; urosomite 3 protuberant dorsally. Rami
of uropods 1-2 with articulate enlarged apical
nails, rami of uropods 1-2 lacking accessory nails,
outer ramus of uropod 1 with 7-12 dorsal spines,
inner with 1-2, outer ramus of uropod 2 with 4-7
dorsal spines, inner with one dorsomedial spine;
peduncle of uropod 1 with 5-6 apicolateral spines
and 0-2 basofacial setae, medially with 6-9 mar-
ginal spines, apicalmost or 2 enlarged; peduncle
of uropod 2 with 14-18 dorsal spines, medially
with one small apical spine. Peduncle of uropod
3 with 6-7 ventral spines, dorsally with one lateral
spine, one medial spine; rami submasculine, inner
extending to M. 95 on article 1 of outer ramus,
apex with 2 setae, medial and lateral margins se-
tose, article 2 of outer ramus short, 0.12, bearing 2
long setae, apicomedial margin of article 1 with
setae, lateral margin with 4 acclivities, spine for-
mula = 1-1-1-1-1, setal formula = 1-1-2-2-1. Tel-
son ordinary, length-width ratio = 11:10, not fully
cleft, each apex wide, rounded, lateral acclivity
shallow, weak, bearing ordinary lateral setule. spine
next medial shorter than setule, midlaicial setules
weakly diverse. Cuticle with sparse ordinary bul-
bar setules.
OBSERVATION (female).?Female "g," length un-
known, damaged, peduncle of uropod 2 with 18
spines.
DESCRIPTION OF MALE.?Rostrum less sharply
constricted than in female. Article 2 of antenna 1
with 4 ventral setae; primary flagellum with 12
articles, one calceolus each on articles 1-6, aes-
thetascs strongly developed. Facial spine formula
on article 4 of antenna 2 = 2-1-4-4, on article
5 = 3 or 2-2 or 2-1; article 5 with 3 sets of dorsal
male setae and one calceolus, ventrodistal apex
with 2 thin spines and one setule, flagellar for-
mula = 35, 2, 3, 5, 7 . . . 34. Right lacinia mobilis
with poorly developed distal branch; palp article 2
of mandible with 2 long apical setae, pair of tiny
basal setules, intervening setae longer than in fe-
male, basofacial setal formula of article 3 = 2 + 3.
Palp of maxilla 1 with 12-13 setae and spines. Palp
article 3 of maxilliped with 1-2 lateral setae. Coxa
4 not broadened; ventral setal formula of coxae
1-4 = 8-8-7-0. Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 = (1-2)-
(3-5)-3-(3-4). Article 6 of pereopods 3-4 slightly
narrowed. Posterior setae of coxae 5-7 shortened.
Epimera 2-3 broadened, posterior margin of epi-
meron 3 not shortened; setal formulas, epimeron
1 anteroventral = 6-8, posteroventral = 2, epimeron
2 facial = 8-9, epimeron 3 posterior = 4, facial =
3, ventral = 2 setules. Articulation of urosomite 1
obscure proximally. Spine formulas of uropods,
uropod 1 peduncle apicolateral = 6, basofacial =
NUMBER 245 S23
nGlt
FIGURE 157.?Birubius kokorus, new species, female "n," 8.2 mm (q ? male "q," 6.3 mm).
324 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 158.?Birubius kokorus, new species, holotype, female "a," 7.7 mm (n ? female "n," 8.2
mm; q = male "q," 6.3 mm; u = female "u," 15 mm).
NUMBER 245 325
1-2, uropod 2 peduncle dorsal = 13-15, dorsal
spines on outer ramus of uropod 1 = 7-8, of uro-
pod 2 = 6, inner ramus of uropod 1 = 1-2, of
uropod 2 = 1 vestigial, ventral spines on peduncle
of uropod 3 = 7, spine formula on article 1 of
outer ramus = 0-1-1-2-2-1, setal formula = 1-1-
1-1-1-1. Telson not elongate, not broadened, distal
spines not shortened.
ILLUSTRATIONS.?Female prebuccal complex and
antenna 1 closely resembling those of male (illus-
trated); lower lip as shown for B. gelarus (17) and
maxilla 2 as shown for female of B. myallus (4).
Male prebuccal complex, maxillae, mandible (ex-
cept for palp and right Iacinia mobilis, separately
illustrated), maxilliped, coxae 2 and 3, gnathopods
and pereopods I, 2, and 3 resembling those of the
female; male pereopod 4 represented in outline
only to show proportions of articles for comparison
with those of female. Drawing of left molar of
female "y" showing apical spines further flattened
in plane of their natural inclination, across apex,
but plume flattened outwards; left maxillipedal
palp of female "n" figured to show those setae nor-
mally hidden or obscured in right aspect as illus-
trated from other side of same specimen; medial
peduncular spinules of uropod 1 and proximoven-
tral spinule on inner ramus not visible in normal
lateral view of urosome; uropod 1 with 0-1 baso-
facial seta on holotype, shown as absent on illus-
tration.
HOLOTYPE.?NMV, female "a," 7.7 mm.
TYPE-LOCALITY.?CPBS 32S/870, 12 Aug 1970,
Western Port, Victoria, Australia, 13 m, muddy
sand, gravel, some shell.
VOUCHER MATERIAL.?Illustrated: Type-locality:
female "n," 8.2 mm. CPBS 32S/269: male "q," 6.3
mm; subadult female "u," 7.5 mm. CPBS 41S/5:
female "y," 7.8 mm.
Not Illustrated: CPBS 41N/4: male "f," 6.1
mm. CPBS 51N/3: female "g," unmeasured, see
observations.
RELATIONSHIP.?This species is difficult to distin-
guish from the B. gambodeni group (20-21) quali-
tatively except in the tightly appressed branches
of the right Iacinia mobilis, and in the presence
of spine(s) on posteroproximal article 5 of pereo-
pods 1-2; adults similar in size to those of B. gam-
bodeni have only 0-2, usually 0-1 basofacial setae
on uropod 1 and can usually be sorted away from
the B. gambodeni group on that basis before the
right mandible must be checked. Full adults of B.
gambodeni have accessory posterior setae on the
telson.
Spines on the telson are shortened in B. kokorus.
Birubius maamus (21) differs from B. kokorus
in the more proximally placed ventral cluster of
setae on article 2 of antenna 1 and also in the
basofacial setae on uropod 1 and the long anterior
ridge on pereopod 5 which in B. maamus are like
those of B. gambodeni.
Birubius kokorus and species 26-27 differ from
species 21-24 in the absence of long ventral setae
on epimeron 3.
MATERIAL.?CPBS, 71 samples from 23 stations
(179); WPBES, 5 samples from 4 stations (9);
WAM, one sample (1).
DISTRIBUTION.?Victoria, Western Port, 2-19 m,
sand, silty sand; New South Wales, off Eden, 30 m,
coarse sand.
26. Birubius kinkus, new species
FIGURES 159, 160
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total body length, greatest width about 60
percent of length; rostrum almost unconstricted,
broad, reaching apex of article 1 on antenna 1.
Eyes medium, largely occluded with pigment. Arti-
cle 1 on peduncle of antenna 1 about 1.4 times
as long as wide, about 2.2 times as wide as article
2, ventral margin with about 10 setules, weakly
produced dorsal apex with 4 setules; article 2 about
0.7 times as long as article 1, with apicoventral
cycle of 6 setae; primary flagellum with 13-15
articles, about 1.2 times as long as peduncle, bear-
ing several short aesthetascs; accessory flagellum
with 10-12 articles. Spine formula on article 4 of
antenna 2 = 1-3-4-4, dorsal margin with notch
bearing 3 setae, ventral margin with 4 groups of
2 long to medium setae, one ventrodistal long
spine; article 5 about 0.7 times as long as article 4,
facial spine formula = 1-2, dorsal margin naked,
ventral margin with 2 sets of 2-3 long to short
setae, 2 ventrodistal long to medium spines; flagel-
lum about 1.8 times as long as articles 4-5 of pe-
duncle combined, with 17 articles. Mandibles with
medium palpar hump; right incisor with 3 teeth;
left incisor with 3 teeth in 2 branches; right Iacinia
mobilis bifid, distal branch much shorter than
326 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
yEU
FIGURE 159.?Birubius kinkus, new species (x ? juvenile "x," 6.0 mm; y - juvenile "y," 5.8 mm).
NUMBER 245 327
proximal, broad, almost subbifid, proximal branch
simple, pointed, with marginal denticles; left la-
cinia mobilis with 4 teeth; right rakers 8; left
rakers 8 plus one rudimentary; molar in form of
elongate plaque, each molar with 8 primarily long
spines, one short very weak spine strongly disjunct,
each molar with plume; palp article 1 short, article
2 with one medium inner apical seta and 3 other
shorter inner setae, article 3 about 0.9 times as
long as article 2, oblique apex with 6 spine-setae,
basofacial formula =1-2 or 1-3. Inner plate of
maxilla 1 ordinary, bearing one long apical plu-
seta, one shorter similar apicomedial seta, 2 apico-
lateral much shorter setae; palp article 2 with 4
apical medial marginal spines and 4 submarginal
setae. Plates of, maxilla 2 extending subequally,
outer as broad as inner, outer with 4-5 apicolateral
setae, inner with one medial seta. Inner plate of
maxilliped with one large thick apical spine, 3
apicofacial setae, 4 medial setae; outer plate with
7 medial and apical spines, 2-3 apicolateral setae;
palp article 1 with apicolateral seta, article 2 with
2 groups of 1-2 apicolateral setae, medial margin
of article 2 moderately setose, article 3 protuberant,
with 5 facial setae, 2 lateral setae, nail of article 4
long-medium, with 2 accessory setules, first of these
0.66 as long as primary nail. Coxa 1 strongly ex-
panded distally, anterior margin almost straight;
main ventral setae of coxae 1-4 = (7-8)-(7-8)-
7-0, posteriormost seta of coxae 1-3 slightly short-
ened; anterior and posterior margins of coxa 4
strongly divergent, posterior margin oblique, al-
most straight, posterodorsal corner rounded, pos-
terodorsal margin short, ordinary, width-length
ratio of coxa 4 = 16:17. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
3-5-5-(5-6), long anteriors = 4-(6-7)-0-O, short
anteriors = 4-5-7-6, no others. Gnathopods ordi-
nary; width ratios of articles 5-6 on gnathopods
1-2 = 22:31 and 22:33, length ratios = 73:60
and 66:55; palmar humps ordinary, palms oblique;
article 5 of gnathopod 1 elongate, ovate, posterior
margin rounded-flat; article 5 of gnathopod 2
elongate, ovate, posterior margin rounded. Pereo-
pods 1-2 similar; facial setae formula on article
4 = 3 and 3, on article 5 = 4 and 5; main spine of
article 5 extending to M. 80-100 on article 6, arti-
cle 5 with 3 and 2 proximoposterior spines; spine
formula of article 6 = 3 + 5 or 4 + 5 plus middis-
tal seta; acclivity on inner margin of dactyls of
pereopods 1-2 weak but produced, emergent setule
short, midfacial pluseta ordinary. Coxae 5-7 pos-
teroventral setule formula = 2-4-3. Articles 4-5 of
pereopods 3-4 of ordinary width, facial spine rows
moderately developed, facial ridge formula on arti-
cle 2 of pereopods 3-5 = 0-2-2; width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 45:47:43:18,
of pereopod 4 = 73:45:34:15, of pereopod 5 =
91:26:20:10, length ratios of pereopod 3 = 78:
33:4450, of pereopod 4 = 96:58:45:54, of pe-
reopod 5 = 124:33:30:32; article 2 of pereopod
5 reaching middle of article 5, one ventral setule
elongate; medial apex of article 6 not combed,
bearing 5 vestigial digital processes. Posteroventral
corner of epimeron 1 rounded, posterior margin
almost straight, corner with setule, anteroventral
margin with 5-7 short setae, posteroventral face
with 2-3 medium setae; posteroventral corner of
epimeron 2 rounded, strongly protuberant, guarded
by setule sinus, posterior margin almost straight,
undulant, facial setae = 6-8 crowded, posteriormost
pair set vertically; posteroventral corner of epi-
meron 3 weakly protuberant, with setule sinus
above and 2 below corner, with small tooth, pos-
terior margin weakly sinuous, with 1-2 setule
notches and 2 long setae, ventral margin naked,
face with horizontal row of 3-4 setae; epimeron
3 with small setule on posterodorsal margin set in
weak notch. Urosomite 1 naked ventrally, articula-
tion line short, almost absent; urosomite 3 pro-
tuberant dorsally. Rami of uropods 1-2 with articu-
late enlarged apical nails, outer ramus of uropod 1
with 5-8 dorsal spines, inner with 1-2, outer ramus
of uropod 2 with 3^1 dorsal spines, inner with one
dorsomedial spine; peduncle of uropod 1 with 5-6
apicolateral spines and 1-2 basofacial setae; medi-
ally with 5 marginal setae and spines, apicalmost
an enlarged spine; peduncle of uropod 2 with 7-12
dorsal spines, medially with one small apical spine.
Peduncle of uropod 3 with 6-7 ventral spines, dor-
sally with one lateral spine, one medial spine and
setule; rami submasculine, inner extending to M.
85 on article 1 of outer ramus, apex with 2 setae,
medial and lateral margins setose, article 2 of outer
ramus short, 0.15, bearing 2 medium setae, medial
margin of article 1 setose but not apically, lat-
eral margin with 4 acclivities, spine formula =
2-2-2-2-2, setal formula = 0-1-1-1-1. Telson long,
length-width ratio = 15:13, not fully cleft, each
apex of medium width, rounded, lateral acclivity
328 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 160.?Birubius kinkus, new species, holotype, female "a," 6.84 mm (x ?? juvenile "x,
6.0 mm; y ? juvenile "y," 5.8 mm).
NUMBER 245 329
deep, narrow, bearing long lateral setule, spine
next medial of length equal to setule, midlateral
setules diverse. Cuticle with ordinary bulbar setules
of varying sizes closely mixed with pipes, setules
bearing apical halo, surface bearing extremely fine
striations in form of linear fingerprint pattern,
emergent setules especially short.
OBSERVATIONS.?Illustrations mainly taken from
juvenile "x," 6.0 mm, noted below; eyes of adult
female slightly enlarged as shown in illustration
from holotype female "a"; uropod 3 of adults as
in B. kokorus (25) but with differences noted in
description above.
Juvenile "x," 6.0 mm, WPBES 51S/1: Eyes clear;
head shorter than in adult; disjunct spine on left
molar broken off (socket present), apparently ab-
sent on right molar; article 5 of pereopods 1-2
with 2 or 1 proximoposterior spines; main spine
distally longer than in adult; article 5 on pereopod
4 slightly shorter than in adult; uropod 1 with only
one basofacial seta (not shown on illustration).
Juvenile "y," 5.8 mm: Uropod 1 lacking baso-
facial seta.
ILLUSTRATIONS.?Eye on lateral view of head
slightly shortened dorsoventrally because of ocular
shrinkage (inward pulling of eyes); maxilla 1 like
that of B. maamus (21), inner plate with 2 small,
one medium, one large seta, outer plate with one
thickened spine, palp with 4 spines and 4 setae;
maxilla 2 like that of B. jirrandus (30) but lobes
slightly broader and inner plate with only one
medial seta; maxilliped like that of B. kokorus
(25) but main spine on inner plate illustrated,
outer plate with 2 lateral setae, face of palp article
3 with 4 setae in tandem, article 3 produced as in
B. kokorus, lateral margin of article 3 with one
seta, palp article 4 illustrated; coxae 1-3 ordinary,
as in B. kokorus.
HOLOTYPE.?NMV, female "a," 6.84 mm.
TYPE-LOCALITY.?CPBS 51 S (resampled), 16 Feb
1972, Western Port, Victoria, Australia, 15.1 m,
coarse sand, anchor in clay.
VOUCHER MATERIAL.?CPBS 51 S/l (sample prior
to 16 Feb 1972): juvenile "x," 6.00 mm (illus.);
juvenile "y," 5.80 mm (illus.).
RELATIONSHIP.?Birubius kinkus differs from B.
gambodeni (20) in the presence of strong postero-
proximal spines on article 5 of pereopods 1-2 and
in the elongation of article 5 on gnathopods 1-2.
From B. maamus (21) to B. kyeemus (23), this spe-
cies differs in the absence of one or more ventral
setae on epimeron 3. From B. kokorus (25) B.
kinkus differs in the presence of 2-3 posteroventral
setae on epimeron 1 while having fewer setae on
epimeron 3, in the more elongate article 5 of the
gnathopods, and in the presence of only one spine
on the iner ramus of uropod 1.
This species is placed in both keys, A and B. It
is placed in Key A because the distal branch of the
right lacinia mobilis is like that of B. gambodeni
in being slightly thickened if not weakly subbifid.
MATERIAL.?CPBS, 4 samples from one station
(9).
DISTRIBUTION.?Victoria, Western Port, 15 m,
coarse sand on clay.
27. Birubius munggai, new species
FIGURES 161-163
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total body length, greatest width about 55
percent of length; rostrum constricted, broad,
reaching middle of article 2 on antenna 1. Eyes
medium, largely occluded with pigment. Article I
on peduncle of antenna 1 about 1.5 times as long
as wide, about 1.7 times as wide as article 2, ven-
tral margin with about 12 setules, weakly produced
dorsal apex with 4 setules; article 2 about 0.7 times
as long as article 1, with ventral crescent of 7
setae; primary flagellum with 13 articles, about
1.15 times as long as peduncle, bearing short
aesthetascs; accessory flagellum with 10 articles.
Spine formula on article 4 of antenna 2 = 1-3-5-5,
dorsal margin with notch bearing 3 setae, ventral
margin with 3 groups of 2 long to medium setae,
one ventrodistal long spine; article 5 about 0.65
times as long as article 4, facial spine formula =
1^ 4, dorsal margin bearing several setae, ventral
margin with 2 sets of 2-3 long to short setae, one
long distal spine next to first main facial spine;
flagellum about 1.65 times as long as articles 4-5
of peduncle combined, with 14 articles. Mandibles
with medium palpar hump; right incisor with 3
teeth; left incisor with 4 teeth in 2 branches; right
lacinia mobilis bifid, distal branch much shorter
than proximal, narrow, simple, proximal branch
simple, pointed, with marginal denticles; left la-
cinia mobilis with 4 teeth; right rakers 9; left rak-
ers 8 plus one rudimentary; molar in form of elon-
330
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 161.?Birubius munggai, new species, holotype, female "a," 6.40 mm (b ? male "b,"
5.30 mm; c ? female "c," 5.40 mm).
NUMBER 245 331
gate plaque, each molar with 7 primarily long
spines, none disjunct, each molar with plume; palp
article 1 slightly elongate, article 2 with 2 medium
inner apical setae and 2 other shorter inner setae,
article 3 about as long as article 2, oblique apex
with 8 spine-setae, basofacial formula = 0-1. Inner
plate of maxilla 1 ordinary, bearing one long apical
pluseta, one shorter similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
2 apicalmedial marginal spines and 4 submarginal
setae. Plates of maxilla 2 extending subequally,
outer scarcely broader than inner, outer with 4
apicolateral setae, inner with 2-3 medial setae. In-
ner plate of maxilliped with one large thick apical
spine, 2 apicofacial setae, 5 medial setae, outer
plate with 7 medial and apical spines, one apico-
lateral seta; palp article 1 with apicolateral seta,
article 2 with one apicolateral seta, medial margin
of article 2 strongly setose, article 3 protuberant,
with 4 facial setae, 2 lateral setae, nail of article 4
long to medium, with 2 accessory setules, one of
these 0.75 as long as nail. Coxa 1 strongly expanded
distally, anterior margin straight; main ventral
setae of coxae 1-4 = 6-6-6-0, posteriormost seta of
coxae 1-2 shortest; anterior and posterior margins
of coxa 4 divergent, posterior margin oblique, al-
most straight, posterodorsal corner rounded, pos-
terodorsal margin short, ordinary, width-length
ratio of coxa 4 = 15:17. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
3-3-4-5, long anteriors = 4-5-0-0, short ante-
riors = 7-7-6-9, no others. Gnathopods ordinary;
width ratios of articles 5-6 on gnathopods 1-2 =
25:40 and 27:39, length ratios = 66:65 and
61:62; palmar humps ordinary, small, palms
oblique; article 5 of gnathopod 1 elongate, ovate,
posterior margin rounded-flat; article 5 of gnatho-
pod 2 ovate, posterior margin rounded, weakly lo-
bate. Pereopods 1-2 similar; facial setae formula
on article 4 = 4 and 4, on article 5 = 4 and 4;
main spine of article 5 extending to M. 80 on arti-
cle 6, article 5 with one proximoposterior spine;
spine formula of article 6 = 3 + 4 and 3 + 4 plus
stiff middistal seta; acclivity on inner margin of
dactyls of pereopods 1-2 weak, emergent setule
short, midfacial pluseta ordinary. Coxae 5-7 pos-
teroventral setule formula = 3-4-4. Articles 4-5 of
pereopods 3^1 of ordinary width, facial spine rows
moderately developed, facial ridge formula on arti-
cle 2 of pereopods 3-5 = 0-2-2; width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 47:43:40:17,
of pereopod 4 = 74:46:34.14, of pereopod 5 =
90:27:23:10, length ratios of pereopod 3 = 80:
37:45:48, of pereopod 4 = 98:56:40:55, of pereo-
pod 5 = 128:33:31:36; article 2 of pereopod 5
reaching middle of article 5; medial apex of article
6 finely combed, bearing 4 vestigial digital proc-
esses. Posteroventral corner of epimeron 1 with
small tooth, posterior margin weakly convex, cor-
ner with setule, anteroventral margin with 6 short
setae, posteroventral face with one medium seta;
posteroventral corner of epimeron 2 rounded,
weakly protuberant, guarded by setule sinus, pos-
terior margin weakly convex, undulant, facial
setae = 6 crowded forward, posteriormost pair set
vertically; posteroventral corner of epimeron 3
weakly protuberant, with setule sinus above and
below small tooth bearing setule, posterior margin
straight, with 2 setules on tooth, ventral margin
naked, face with horizontal row of 3 setae and off-
set ventral seta, epimera 2-3 with tiny setule set
in weak notch. Urosomite 1 with 2 ventral setules
at base of uropod 1, articulation line complete;
urosomite 3 protuberant dorsally. Rami of uropods
1-2 with articulate apical nails and weak accessory
setule, outer ramus of uropod I with 7 dorsal
spines, inner with one, outer ramus of uropod 2
with 5 dorsal spines, inner with one dorsomedial
spine; peduncle of uropod 1 with" 4 apicolateral
spines, and one basofacial seta, medially with 6
marginal setae and spines, apicalmost an enlarged
spine; peduncle of uropod 2 with 13 dorsal spines,
medially with one small apical spine. Peduncle of
uropod 3 with 6 ventral spines, dorsally with one
lateral spine, one medial spine and setule; rami
submasculine, inner extending to M. 100 on article
1 of outer ramus, apex with 3 setae, medial and
lateral margins setose, article 2 of outer ramus
short, 0.15, bearing 2 long setae, apicomedial mar-
gin of article 1 setose, lateral margin with 4 ac-
clivities, spine formula = 2-1-1-1-1, setal for-
mula = 0-1-1-2-2. Telson weakly elongate, length-
width ratio = 29:27, not fully cleft, each apex
wide, rounded, lateral acclivity deep, narrow, bear-
ing long lateral setule, spine next medial shorter
than setule, midlateral setules diverse. Cuticle with
ordinary bulbar setules, surface bearing fine stria-
tions in form of linear fingerprint pattern, emerg-
ent setules especially short.
DESCRIPTION OF MALE.?Article 2 of antenna 1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 162.?Birubius munggai, new species, holotype, female "a," 6.40 ram {b ? male "b,1
5.30 mm).
NUMBER 245 333
FIGURE 163.?Birubius munggai, new species, holotype, female "a," 6.40 mm (b ? male "b,"
5.30 mm; c ? female "c," 5.40 mm; y = male "y," 5.50 mm).
with numerous ventral setules; primary flagellum
with 13 articles, one calceolus each on articles 1-5;
accessory flagellum with 11 articles. Facial spine
formula on article 4 of antenna 2 = 3-5-5; article
5 with 3 dorsal sets of male setae and one calceolus,
ventrodistal apex with 2 thin spines and one setule;
flagellar formula = 31, 2, 3, 5, 7 . . . 29. Right la-
cinia mobilis with distal branch shorter than in
female; left rakers 9 plus 1-2 rudimentaries; molars
with 6 spines; basofacial setae on palp article 3 =
2 + 2. Coxa 4 broadened but smaller in relation
to coxa 1 than in female. Article 2 of pereopods
3-5 narrower than in female. Epimera 1-3 broad-
ened, posterior margin of epimeron 3 not short-
ened, convex; setal formulas, epimeron 1 antero-
ventral = 5, posteroventral = 1, epimeron 2 facial =
6, epimeron 3 posterior = 2 long, facial = 3 setae,
ventral = 0. Spine formulas of uropods, uropod 1
334 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
peduncle apicolateral = 5, basofacial = 1, uropod
2 peduncle dorsal = 13, dorsal spines on outer ra-
mus of uropod 1 = 6, of uropod 2 = 5, inner ramus
of uropod 1 = 1, of uropod 2 = 1, ventral spines
on peduncle of uropod 3 = 6, spine formula on
article 1 of outer ramus = 0-1-1-1-1-1-1, setal for-
mula = 1-1-0-1-1-1-1. Telson weakly elongate,
distal spines scarcely shortened.
OBSERVATIONS.?Female "c," 5.40 mm, and male
"y," 5.50 mm, lacking basofacial seta on uropod 1.
Telson of female "c" aberrant, one side lacking
spine.
ILLUSTRATIONS.?Following parts resembling Biru-
bius kokorus (25) except as specified in description:
prebuccal parts, upper lip, mandibles, maxilla 1,
maxilliped, dactyls of pereopods 1-2.
HOLOTYPE.?NMV, female "a," 6.40 mm.
TYPE-LOCALITY.?PPBES 126, 9 Mar 1971, Port
Phillip Bay, Victoria, Australia, 8 m, sand.
VOUCHER MATERIAL (all illustrated).?PPBES
986/5: male "b," 5.30 mm; female "c," 5.40 mm;
male "y," 5.50 mm.
RELATIONSHIP.?This species is extremely close
to and may be a phenotype of Birubius kokorus
(25) but differs in the slightly elongate article 1
of the mandibular palp, the presence in adults of
only one dorsal spine on the inner ramus of uropod
1 and in the female by the fewer ventral sets of
setae on article 5 of antenna 2. Birubius munggai
bears only one posteroproximal spine on article 5
of pereopods 1-2.
From B. kyeemus (23) and B. lowannus (22) this
species differs in the elongate palp article 1 of the
mandible and in the absence of the one ventral
seta on epimeron 3 known for B. kyeemus and B.
lowannus.
Birubius kinkus (26) differs from B. munggai in
the shorter article 1 of the mandibular palp, the
consistently more elongate wrists of the gnatho-
pods, the more strongly setose epimeron 1, and the
presence of more than one posteroproximal spine
on article 5 of pereopods 1-2. Again, B. munggai
may be a phenotype of either B. kokorus or B.
kinkus but the elongation of mandibular palp arti-
cle 1 occurs well into adulthood and is not found
in the other two species.
MATERIAL.?CPBS, one sample (1); PPBES, 2
samples from 2 stations (5).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 4-13 m, sand, sandy gravel.
28. Birubius ularitus, new species
FIGURES 164-166
DESCRIPTION OF MALE (subadult).?Head about
21 percent of total body length, greatest width
about 67 percent of length; rostrum unconstricted,
broad, exceeding middle of article 2 on antenna 1.
Eyes medium, clear of pigment core but deeply
burgundy. Article 1 on peduncle of antenna 1 about
1 25 times as long as wide, about twice as wide as
article 2, ventral margin with about 10 setules,
weakly produced dorsal apex with 3 setules; article
2 about 0.7 times as long as article 1, with ventral
cycle of 5 setae; primary flagellum with 11 articles,
about 0.9 times as long as peduncle, bearing long
aesthetascs; accessory flagellum with 8 articles. Spine
formula on article 1 of antenna 2 1-3-4? i, dorsal
margin with notch bearing 4 setae, ventral margin
with 7 groups of 1-2 long to short setae, one ven-
trodistal long spine; article 5 almost 0.75 times as
long as article 4, facial spine formula = 1-2-2,
dorsal margin bearing one set of male setae, ventral
margin with 4 sets of 2 long to short setae, 2 ventro-
distal long to medium spines, one set facially; flagel-
lum proliferate as young male. Mandibles with
weak palpar hump; right incisor with 3 teeth; left
incisor with 4 humps in 2 branches; right lacinia
mobilis simple, denticulate; left lacinia mobilis with
4 teeth plus one accessory tooth; right rakers 8; left
rakers 9 plus one rudimentary; molar in form of
elongate bulbous hump, demarcated mainly by
spines, right molar with 7 primarily long spines
plus one setule, none disjunct, left molar with 7
primarily long spines, only left molar with plume;
palp article 1 short, article 2 with one medium
inner apical seta and 2 other shorter inner setae,
article 3 about 1.25 times as long as article 2,
oblique apex with 8-9 spine-setae, basofacial
formula = 1-2 or 1-1. Inner plate of maxilla 1
large, thin, bearing one long apical pluseta, one
shorter similar apicomedial seta, 2 apicolateral
much shorter setae; palp article 2 with 4 apicalmed-
ial marginal spines and 4 submarginal setae. Plates
of maxilla 2 extending equally, of subequal width,
outer with 3 apicolateral setae, inner with one
medial seta. Inner plate of maxilliped with 2 large,
thick apical spines, 2 apicofacial setae, 4 medial
setae, outer plate with 10 medial and apical spines,
4 apicolateral setae; palp article 1 with apicolateral
seta, article 2 with one apicolateral seta, medial
NUMBER 245 335
FIGURE 164.?Birubius ularitus, new species, holotype, male "y," 4.2 mm.
margin of article 2 weakly setose, article 3 with 4
facial setae, one lateral seta, nail of article 4 med-
ium, with 2 accessory setules. Coxa 1 scarcely ex-
panded distally, anterior margin almost straight;
main ventral setae of coxae 1-4 = 8-7-6-11, pos-
teriormost seta of coxae 1-4 shortened, penultimate
seta especially stout on coxae 1-3; anterior and
posterior margins of coxa 4 strongly divergent,
posterior margin oblique, almost straight, postero-
dorsal corner sharp to rounded, posterodorsal mar-
gin short, weakly concave, width length ratio of
coxa 4 = 16:17. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 2-2-2?4, long
anteriors = 4(4- 1 facial)-6-0-0, short anteriors =
1-1-3-2, no others. Gnathopods ordinary; width
ratios of articles 5-6 on gnathopods 1-2 = 24:33
and 26:34, length ratios = 60:60 and 55:57, palmar
humps ordinary to large, palms strongly oblique;
336
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
PS
FIGURE 165.?Birubius ularitus, new species, holotype, male "y," 4.2 mm.
article 5 of gnathopod 1 weakly elongate, ovate,
posterior margin rounded; article 5 of gnathopod 2
ovate, posterior margin rounded. Pereopods 1-2
similar; facial setae formula on article 4 = 6 and 6,
on article 5 = 6 and 6; main spine of article 5 ex-
tending to M. 85 on article 6, article 5 lacking
proximoposterior spines; spine formula of article
6 = 3 + 4 and 4 + 4 plus middistal seta; acclivity
on inner margin of dactyls of pereopods 1-2 weak,
emergent setule short, midfacial pluseta ordinary.
Coxae 5-7 posteroventral setule formula = 5-3-1.
Articles 4?5 of pereopods 3-4 narrow to ordinary in
width, facial spine rows moderately developed,
facial ridge formula on article 2 of pereopods 3-5
= 0-2-2; width ratios of articles 2, 4, 5, 6 of pereo-
pod 3 = 37:32:27:14, of pereopod 4 = 55:26:20:10,
of pereopod 5 = 76:20:18:6, length ratios of
pereopod 3 = 63:30:33:36, of pereopod 4 = 80:
45:38:54, of pereopod 5 = 98:25:24:27; article 2
of pereopod 5 not reaching apex of article 4,
with one ventral seta; medial apex of article 6
scarcely combed, bearing 5 digital processes. Postero-
NUMBER 245 337
FIGURE 166.?Birubius ularitus, new species, holotype, male "y," 42 mm.
ventral corner of epimeron 1 weakly protuberant,
posterior margin convex, with one setule, corner
with setule, anteroventral margin with 4 medium
setae, posteroventral face with 2 long setae; postero-
ventral corner of epimeron 2 weakly protuberant,
with setule sinus, posterior margin convex, with one
setule, facial setae =10, posteriormost pair set
vertically; posteroventral corner of epimeron 3
weakly protuberant, with setule sinus, with small
tooth, posterior margin weakly convex, with 2 setule
notches, mid ventral margin with 3 setae, epimera
1-3 with small seta on posterodorsal margin set in
weak to strong notch. Urosomite 1 with lateral
setule at base of uropod I, articulation line almost
complete; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1-2 with articulate enlarged
apical nails, outer ramus of uropod 1 with 5 dorsal
spines, inner with 2, outer ramus of uropod 2 with
3 dorsal spines, inner with one dorsomedial spine;
peduncle of uropod 1 with 2 apicolateral spines and
5 basofacial setae, medially with 4 thin marginal
spines plus apical enlarged spine; peduncle of uro-
pod 2 with 8 dorsal spines, medially with one small
apical spine, peduncle of uropod 3 with 4 ventral
spines, dorsally with 2 lateral spines or spine and
setule, one medial setule; rami submasculine, inner
extending to M. 95 on article 1 of outer ramus, apex
with 2 short setae, medial margin with one seta,
article 2 of outer ramus short, 0.13, bearing 2 long
setae, apicomedial margin of article 1 with 2 setae,
lateral margin with 4 acclivities, spine formula =
1-1-1-1-1, setal formula = 0-1-1-2-2. Telson long,
length-width ratio =13:11, not fully cleft, each
apex wide, rounded, lateral acclivity broad, shallow,
bearing ordinary lateral setule, spine next medial of
length equal to setule, midlateral setules diverse.
Cuticle with bulbar setules of varying sizes mixed
with pipes, surface bearing fine striations in form of
linear fingerprint pattern, emergent setules ordi-
nary.
ILLUSTRATION.?Uropod 3 forming composite of
right and left members to show maximum number
of setae and spines in each position.
HOLOTYPE.?NMV, young male "y," 4.2 mm.
Unique.
TYPE-LOCALITY.?CPBS A2/3, 15 Jul 1964, West-
ern Port, Victoria, Australia, 13.5 m, sediment un-
known.
RELATIONSHIP.?This species and B. eleebanus
(29) may be distinct generically from Birubius.
Pereopod 4 is very thin and the right lacinia
mobilis may actually be totally absent so that what
appears to be the right lacinia mobilis is really the
first raker spine slightly modified; the odd base of
the spine suggests this interpretation. This possibil-
ity should be studied embryologically. The thin
338 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pereopod 4, however, is matched in other species of
Birubius such as B. mayamayi (11).
In other respects this species is almost identical
to B. mayamayi but differs in the simple right
lacinia mobilis. Pereopod 5 bears one ventral seta
on article 2 thereby suggesting relationship to B.
karobrani (14) but again the simple right lacinia
mobilis distinguishes B. ularitus. The similarities of
B. ularitus to Cunmurra itickerus are striking.
Birubius ularitus might form the ancestral condi-
tion for B. wulgaru (37). The only species with sim-
ple right lacinia mobilis to which B. ularitus bears
comparison is B. eleebanus; the latter lacks the
extra serrations on epimeron 3 and has a more com-
plex cuticle.
MATERIAL.?CPBS, one sample (1).
DISTRIBUTION.?Victoria, Western Port, 13.5 m.
29. Birubius eleebanus, new species
FIGURES 167-170
DESCRIPTION OF FEMALE.?Head about 19 per-
cent of total body length, greatest width about 75
percent of length; rostrum slightly constricted,
short, reaching apex of article 1 on antenna 1. Eyes
medium, clear of pigment, ommatidia especially
large. Article 1 on peduncle of antenna 1 about 1.3
times as long as wide, about 2.1 times as wide as
article 2, ventral margin with about 8 setules, un-
produced dorsal apex with 4 setules; article 2
about 0.9 times as long as article 1, with apicoven-
tral cycle of 4-5 setae; primary flagellum with 8
articles, about 0.87 times as long as peduncle, bear-
ing aesthetascs; accessory flagellum with 5-6 arti-
cles. Spine formula on article 4 of antenna 2 =
1-3-4-4, dorsal margin bearing 2 setae, ventral
margin with 4 groups of 1-2 long to medium setae,
one ventrodistal long spine; article 5 about 0.9
times as long as article 4, facial spine formula =
1-2-2, dorsal margin naked, ventral margin with 3
sets of 1-2 long to medium setae, 2 ventrodistal me-
dium spines; flagellum about 1.15 times as long as
articles 4-5 of peduncle combined, with 8-9 arti-
cles. Mandibles with weak palpar hump; right in-
cisor with 3 teeth and notch; left incisor with 4
humps in 2 branches; right lacinia mobilis simple,
denticulate; left lacinia mobilis with 4 teeth, mid-
dle teeth shortened; right rakers 8 plus one rudi-
mentary; left rakers 8; molar in form of elongate
plaque, molars with 8-9 primarily long spines,
none disjunct, each molar with plume; palp article
1 slightly elongate, article 2 with one long inner
apical seta and 2 other short inner setae, article 3
about 1.1 times as long as article 2, oblique apex
with 6 spine-setae, basofacial formula = 1-1. Inner
plate of maxilla 1 large, bearing one long apical
pluseta, one shorter apicomedial seta, 2 apicolateral
much shorter setae; palp article 2 with 2 apical-
medial marginal spines and 4 submarginal setae.
Plates of maxilla 2 extending equally, inner slightly
broader than outer, outer with 3 apicolateral setae,
inner with 2 medial setae. Inner plate of maxilliped
with 2 large, thick apical spines, 2 apicofacial setae,
3 medial setae; outer plate with 6 medial and apical
spines, 2 apicolateral setae; palp articles 1-2 with
apicolateral seta, medial margin of article 2 weakly
setose, article 3 unprotubcrant with 3 facial setae,
no lateral setae, nail of article 4 medium, with 2
accessory setules. Coxa 1 strongly expanded distally,
anterior margin weakly convex; main ventral setae
of coxae I--1 = (7-12>-<8-10)-(9-l l>-<S-5), poste"
riormost seta of coxae 1-3 shortened, one middle
seta especially stout; anterior and posterior margins
of coxa 4 strongly divergent, posterior margin very
oblique, almost straight, posterodorsal corner
rounded, posterodorsal margin short, undulant,
width-length ratio of coxa 4 = 1:1. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 2-3-2-4, long anteriors = 3-6-0-0, short
anteriors = 1-1-2-2, no others. Gnathopods with
elongate wrists and broad hands; width ratios of
articles 5-6 on gnathopods 1-2 = 23:40 and 25:40,
length ratios = 74:65 and 65:62; palmar humps
large, palms weakly oblique; article 5 of gnatho-
pods 1-2 elongate, ovate, posterior margin rounded.
Pereopods 1-2 similar, facial setae formula on arti-
cle 4 = 4 and 6, on article 5 = 4 and 5; main spine
of article 5 extending to M. 85 on article 6, article
5 lacking proximoposterior spines; spine formula of
article 6 = 3 + 4 plus middistal seta; acclivity on
inner margin of dactyls of pereopods 1-2 weak,
sharp, produced as tooth, emergent setule long,
midfacial pluseta very long. Coxae 5-7 posteroven-
tral setule formula = 5-3-5. Articles 4-5 of pereo-
pods 3-4 of ordinary width, facial spine rows sparse,
facial ridge formula on article 2 of pereopods 3-5 =
1-2-2; width ratios of articles 2, 4, 5, 6 of pereopod
3 = 41:36:36:20, of pereopod 4 = 60:36:31:12, of
pereopod 5 = 78:23:22:9, length ratios of pereopod
NUMBER 245 339
FIGURE 167.?Birubius eleebanus, new species, holotype, female "a," 3.86 mm (b ?? male "b,"
3.57 mm).
340 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
3 = 66:33:37:9, of pereopod 4 = 80:52:42:53, of
pereopod 5 = 98:25:26:26; article 2 of pereopod
5 reaching apex of article 4; medial apex of
article 6 finely combed, bearing obsolescent digital
processes. Posteroventral corner of epimeron 1
weakly and sharply protuberant, posterior margin
almost straight, anteroventral margin with 1-2 long
setae, posteroventral face with one long seta; pos-
teroventral corner of epimeron 2 weakly protuber-
ant, posterior margin convex, facial setae = 6; poste-
roventral corner of epimeron 3 weakly protuberant,
with setule sinus, with small tooth, posterior mar-
gin weakly convex, with one large to small dorsal
seta, ventral submargin with 4-5 setae mainly in
middle of margin. Urosomite 1 naked, articulation
line almost complete; urosomite 3 weakly protuber-
ant dorsally. Rami of uropods 1-2 with articulate
enlarged apical nails, outer ramus of uropod 1 with
3 dorsal spines, inner with one, outer ramus of uro-
pod 2 with 1-2 dorsal spines, inner shortened, with
one dorsomedial spine; peduncle of uropod 1 with
2 apicolateral spines and 2-4 basofacial setae, me-
dially with 5 marginal setae and spines, apicalmost
an ordinary spine; peduncle of uropod 2 with 3-4
dorsal spines, medially with one apical spine. Pe-
duncle of uropod 3 with no ventral spines or one
setule only, dorsally with one lateral spine, one
medial spine and one setule; rami feminine, inner
extending to M. 40 on article 1 of outer ramus,
apex with one seta, medial and lateral margins
naked, article 2 of outer ramus short, 0.17, bearing
2 long setae, apicomedial margin of article 1 naked,
lateral margin with 2-3 acclivities, spine formula =
0, setal formula = 1-2-2-2 or 1-2-2. Telson short,
length-width ratio = 5:6, not fully cleft, each
apex wide, rounded, subtruncate, lateral acclivity
broad, shallow, bearing long lateral setule, spine
next medial much shorter than setule, midlateral
setules diverse. Cuticle with occasional bulbar set-
ule set on edge of coronate plaque in midst of
patches of dense polygonal structure formed of
fused scales, surface otherwise covered with flat
scales, many with central points, emergent setules
especially long and branched.
OBSERVATIONS (subadult females).?Midapex of
article 6 on pereopod 5 lacking comb in subadults.
DESCRIPTION OF MALE.?Rostrum smaller than in
female. Eyes largely occluded with pigment. Article
2 of antenna 1 with 4 ventral setae; primary flagel-
lum with 8 articles, one calceolus each on articles
1-3 or 1-6, aesthetascs strongly developed; accessory
flagellum with 5 articles. Facial spine formula on
article 4 of antenna 2 = 0-3-4-4, on article 5 = 2-2,
dorsodistal spine of female absent, article 5 with 3
dorsal sets of male setae and 2 calceoli, ventrodistal
apex with 2 thin spines; flagellar formula = 21, 2-4,
6,8 . . .20. Right rakers 7; left rakers 7 plus one rudi-
mentary; right molar with 6 spines and plume, one
of these spines short but not disjunct, left molar
with 7 spines, one of these short and highly dis-
junct; basofacial setal formula of article 3 on man-
dibular palp = 1-2, terminal = 7-8. Palp of maxilla
1 with 3 spines and 4 setae. Inner plate of
maxilla 2 with one medial seta, outer plate with S
apicolateral seatc. Coxa 4 shaped as in female but
slightly larger in relation to coxa I; ventral setal
formula of coxae 1-4 = 9-8-(7-8)-2. Long poste-
rior setae on article 2 of gnatho|x>ds 1-2 and pereo-
pods 1-2 = 2-2-1-3, long anterior* = 3-5-0-0,
short anteriors = 1-1-1-1, no others. Article 6 of
gnathopods 1-2 slightly larger relative to article 5
than in female. Facial and setal spine formulas of
pereopods 1-2 on article 4 = 4 and 4, on article
5 = 5 and 5, on article 6 = 3 + 4 plus middistal
seta. Article 2 of pereopods 3-5 narrower than in
female, article 5 of pereopod 5 with special spine.
Epimera 1-3 broadened, posterior margin of epi-
meron 3 shortened; setal formulas, epimeron 1 an-
teroventral = 6, posteroventral = 1, posterior =
1 + 1, epimeron 2 facial = 6 + 1 {aberrant), pos-
terior = 1 + 1, epimeron 3 posterior = 1 + 1, fa-
cial = 6. Spine formulas of uropods, uropod 1 pe-
duncle apicolateral = 2, basofacial = 5, uropod 2
peduncle dorsal = 7, dorsal spines on outer ramus
of uropod 1 = 4, of uropod 2 = 2, inner ramus of
uropod 1 = 1, of uropod 2 = 1 . Urosomite 1 with
ventral seta. Ventral spines on peduncle of uropod
3 = 0 , spine formula on article 1 of outer ramus =
1?1?1?1?1?1?1, long setal formula = 1?1?1?1?1?
1-1, short setal formula = 0-0?1?I?1?1-1. Telson
elongate, distal spines shortened. Scales of cuticle
as dense as on female but more evenly concen-
trated, polygons present but not as strongly, out-
lined, bare coronate plaques poorly developed.
ILLUSTRATION.?Female coxa 5 with one setule
missing.
HOLOTYPE.?The figures herein presented of fe-
male "a," 3.86 mm. (Holotype designated as figures
because of possible confoundment of holotype spe-
cimen; senior author accidentally mixed together
NUMBER 245 341
FIGURE 168.?Birubius eleebanus, new species, holotype, female "a," 3.86 mm (b = male "b,"
3.57 mm).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
R2i
FIGURE 169.?Birubius eleebanus, new species, holotype, female "a," 3.86 mm (b ? male "b"
3.57 mm).
NUMBER 245
parts of this holotype plus holotype of B. lorus (2);
in process of segregating these parts telson of this
species lost; one lower lip and one maxilla 2 of this
spedes possibly mixed in with parts of B. lorus.)
TYPE-LOCALITY?SBS D2S2, 17 May 1972, off
Malabar, New South Wales, Australia, 40 m, coarse
sand.
VOUCHER MATERIAL.?SBS 4, male "b," 3.57 mm
(illus.); SBS A2S3, female "c," 3.22; SBS D2S1, fe-
male "d," 3.60 mm.
RELATIONSHIP.?This species differs from B.
ularitus (28), for which only the male is known, in
the shorter and slightly narrowed rostrum, in the
presence of only one notch posteroventally on epi-
meron 3, the much stouter and fewer spines on
male uropod 2, and in the complex cuticle.
Possible affinities of these two species may lie
with the B. babaneekus group (16-18) owing to the
setose coxa 4 but B. ularitus and B. eleebanus have
fully simple and aberrant (see B. ularitus) right
lacinia mobili, small tooth on epimeron 3 and the
presence of a fully dorsoposterior setule on epi-
meron 3, itself suggesting affinity with B. narus (19),
a satellite member of the B. babaneekus group
which also has a large tooth on epimeron 3.
FIGURE 170.?Birubius eleebanus, new species,
female "a," S.86 mm.
holotype.
This species is one of those with numerous web-
like interrelationships. It may be a close sibling of
B. chintoo (13), of which only the male is known.
The resemblance is especially close in shape and
general setation of epimera 1-3, coxa 4, mandibular
molar, telson, uropods 1-2 and gnathopods. Biru-
bius eleebanus differs from B. chintoo in the simple
right lacinia mobilis, broader and shorter head,
extra spines on article 5 of antenna 2, presence of
an anterior ridge on article 2 of pereopod 4, the
presence of a long dorsoposterior seta on epimeron
3 while simultaneously having fewer setae on epi-
mera 1-2, and a greater number of setae on epi-
meron 3. All but one of these characters might be
dismissed if B. chintoo were a supermale, because in
terminal males there is a tendency for reduction of
epimeron 3 setae and reduction of spines on an-
tenna 2 and there may be sexual differences in
facial ridges on pereopods. The crucial distinction
remains the simple right lacinia mobilis of B.
eleebanus, confirmed in the male.
A more distant relationship is seen to B. gal-
langus (10), B. lorus (2) and B. nammuldus (3), all
of which have accessory nails on uropod 1 and a
normal right lacinia mobilis. Birubius karobrani
(14) has ventral setae on pereopod 5 among other
distinctions. The long posterodorsal seta on epi-
meron 3 of B. eleebanus is reminiscent of this con-
dition in the B. mayamayi (ll)-ZJ. karobrani group
but it is more ventrally placed in B. eleebanus than
in the other species. Basically, B. eleebanus could
be termed a member of the B. mayamayi (11) group
in which the right lacinia mobilis has been reduced
to simplicity. See "Remarks" under Parharpinia
villosa.
MATERIAL.?SBS, 4 samples from 3 stations (4).
DISTRIBUTION.?New South Wales, off Malabar,
48-49 m, fine sand.
30. Birubius jirrandus, new species
FIGURES 171-175
DESCRIPTION OF FEMALE.?Head about 20 percent
of total body length, greatest width about 60 per-
cent of length; rostrum weakly constricted, reaching
middle of article 2 on antenna 1. Eyes medium,
clear of pigment. Article 1 on peduncle of antenna
1 about 1.5 times as long as wide, about 2.3 times
as wide as article 2, ventral margin with 9-13
344
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 171.?Birubius jirrandus, new species, holotype, female "a," 9.8 mm (u ? female "u,
9.00 mm; v = male "v," 8.7 mm; y ? male "y," 8.1 mm).
NUMBER 245 345
setules, weakly produced dorsal apex with 3 setules;
article 2 about 0.65 times as long as article 1, with
proximoventral cycle of 5-7 setae; primary flagel-
lum with 10-11 articles, about 0.6 times as long as
peduncle, bearing aesthetascs; accessory flagellum
with 9-10 articles. Spine formula on article 4 of
antenna 2 = 1-3-4-4 or 1-3-3-4, dorsal margin
with notch bearing 3 setae and spines, ventral
margin with 6 groups of 1-3 long to medium setae,
one ventrodistal medium spine; article 5 about 0.9
times as long as article 4, facial spine formula =
1-3 or 1-4, dorsal margin naked, ventral margin
with 4-5 sets of 1-2 long to short setae, 2 ventro-
distal long to medium spines; flagellum about 1.3
times as long as articles 4-5 of peduncle combined,
with 12-14 articles. Mandibles with weak to med-
ium palpar hump; right incisor with 2 teeth; left
incisor with 2 humps in 2 branches; right lacinia
mobilis simple, short; left lacinia mobilis with 5
teeth plus one accessory tooth; right and left rakers
6-8 plus 1-2 rudimentaries; molar in form of short
protrusion demarcated mainly by spines, right
molar with 5-6 primarily long spines plus one spine
strongly disjunct, left molar with 5 primarily long
spines plus one spine strongly disjunct, each
molar with plume; palp article 1 short, article
2 with 2-3 medium inner apical setae and 2-3
other shorter inner setae, article 3 almost 0.95
times as long as article 2, oblique apex with 8-9
spine-setae, basofacial formula = 0-1 or 0-2. Inner
plate of maxilla 1 large, bearing one long apical
pluseta, one shorter similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
4 apicalmedial marginal spines and 4-6 submar-
ginal setae. Plates of maxilla 2 extending equally,
slender, outer not broader than inner, with 4 api-
colateral setae, inner with 2 medial setae. Inner
plate of maxilliped with one large thick apical
spine, 2 apicofacial setae, 4 medial setae; outer plate
short, with 4-6 medial and apical spines, one api-
colateral seta; palp article 1 with 4 apicolateral
setae, article 2 with 4-5 apicolateral setae, medial
margin of article 2 strongly setose, article 3 pro-
tuberant, with 4 proximal facial setae, 2 apicolateral
facial setae, nail of article 4 medium, with 2 acces-
sory setules. Coxa 1 strongly expanded distally,
anterior margin straight; main ventral setae of
coxae 1-4 = 8-9-10-0, posteriormost seta of coxae
1-3 elongate; anterior and posterior margins of coxa
4 strongly divergent, posterior margin very oblique,
convex, posterodorsal corner rounded, posterodorsal
margin short, undulant, width-length ratio of coxa
4 = 15:16. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = (l-4)-(6-7)?
(4-5)-6, and 3 long facials on gnathopod 1, long
anteriors = 7-19-0-0, short anteriors = 2-1-2-2.
Gnathopods generally ordinary but hands thin;
width ratios of articles 5-6 on gnathopods 1-2 =
29:33 and 28:34, length ratios = 70:66 and
63:62; palmar humps ordinary, palms oblique;
article 5 of gnathopod 1 elongate, ovate, posterior
margin flat; article 5 of gnathopod 2 ovate, poste-
rior margin shorter, flat, produced. Pereopods 1-2
similar; facial setae formula on article 4 = 4 and 4,
on article 5 = 5 and 5; main spine of article 5 ex-
tending to M. 80 on article 6, article 5 with 2 proxi-
moposterior spines; spine formula of article 6 =
4 + 4 plus middistal seta; acclivity on inner margin
of dactyls of pereopods 1-2 weak, emergent setule
long, midfacial pluseta ordinary. Coxae 5-7 pos-
teroventral setule formula = (8-9)-4-4. Articles 4-5
of pereopods 3-4 broad to ordinary, facial spine
rows moderately to well developed, facial ridge
formula on article 2 of pereopods 3-5 = 0-2-2,
anterior ridge on pereopod 5 short; width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 37:41:35:17, of
pereopod 4 = 60:37:27:14, of pereopod 5 = 70:
25:22:8, length ratios of pereopod 3 = 66:28:32:
35, of pereopod 4 = 75:41:44:55, of pereopod 5 =
87:27:25:27; article 2 of pereopod 5 almost reaching
apex of article 4; medial apex of article 6 not
finely combed, bearing 6 digital processes. Postero-
ventral corner of epimeron 1 weakly protuberant,
corner with 1-2 setules, anteroventral margin
and face with 5-8 medium setae, posteroventral
face with 3^ 1 long setae, posteriormost pair
occasionally subvertical; posteroventral corner of
epimeron 2 weakly protuberant, with small blunt
sinuous tooth guarded by setule sinus, posterior
margin weakly sinuous, with or without setule,
facial setae = 9-11 crowded forward, posteriormost
pair set vertically; posteroventral corner of epi-
meron 3 weakly protuberant, with setule sinus and
small tooth, posterior margin almost straight, with
4-6 upthrust setae, with 1-2 setule notches, ventral
margin with 2-6 setae, mainly posterior; epimera
1-3 with setule on posterodorsal margin, (missing
on illustration of epimeron 1). Urosomite 1 setose
ventrally at base of uropod 1, articulation line in-
complete, almost absent in full adults, stronger in
346
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 172.?Birubius jirrandus, new species, holotype, female "a," 9.8 mm (v ? male "v," 8.7
mm; y ?? male "y," 8.1 mm).
NUMBER 245 347
subadults; urosomite 3 protuberant dorsally. Rami
of uropods 1-2 with articulate enlarged apical nails,
outer ramus of uropod 1 with 5 dorsal spines, inner
with one, outer ramus of uropod 2 with 3-4 dorsal
spines, inner with one dorsomedial spine; peduncle
of uropod 1 with 3-4 apicolateral spines, and 7-9
basofacial setae, medially with 6 marginal spines,
apicalmost enlarged; peduncle of uropod 2 with
8-10 dorsal spines, medially with one small apical
spine. Peduncle of uropod 3 with 5-6 ventral spines,
dorsally with 1-2 lateral spines, one medial spine
and occasional setule; rami submasculine, inner ex-
tending to M. 80-90 on article 1 of outer ramus,
apex with 2-3 setae, medial and lateral margins
setose, article 2 of outer ramus short, 0.14, bearing
2 long setae, apicomedial margin of article 1 setose,
lateral margin with 4-5 acclivities, spine forumla =
( l ) - l - I - l - l - l , setal formula - 0-l-l-l-(l-2)-2.
Telson weakly elongate, length-width ratio =
24:23, not fully cleft, each apex wide, rounded,
lateral acclivity broad, shallow, bearing ordinary
lateral setule, spine next medial shorter than setule,
midlateral setules diverse. Cuticle with sparse
bulbar setules, emergent setules especially short.
OBSERVATIONS (female).?Article 3 of antenna 2
with short fuzzy crest reminiscent of male. Outer
plate of maxilla 1 with 9 spines.
DESCRIPTION OF MALE.?Rostrum slightly nar-
rower than in female. Primary flagellum of antenna
1 with 11-12 articles, one calceolus each on articles
2-6, accessory flagellum with 9 articles. Facial spine
formula on article 4 of antenna 2 = 3 4 4 or 2-3-4,
on article 5 = 3, article 5 with 2 dorsal sets of male
setae and one calceolus, ventrodistal apex with 2
thin spines and one setule; flagellar formula = 28,
1, 2, 4, 6 . . . 26 or 2, 4, 6 . . . . Basofacial setal for-
mula of article 3 on mandibular palp = 2-1 (same
face), article 2 with group of 3-4 inner apical setae
plus proximal setae. Palp of maxilla 1 with 3 spines
and 4 setae. Outer plate of maxilliped elongate;
palp articles 1-2 with fewer lateral setae than in
female. Coxa 4 broadened but smaller in relation to
coxa 1 than in female; ventral setae of coxae 1-4 =
fewer than in female and deeply inserted. Article 2
of pereopods 3-5 narrower than in female, articles
5-6 of pereopod 4 more attenuate than in female.
Epimera 1-3 broadened, posterior margin of epi-
meron 3 slightly shortened; setal formulas, epi-
meron 1 anteroventral = 7-9, posteroventral = 2-3,
epimeron 2 facial = 7-8, epimeron 3 posterior = 4,
ventral = 2-3. Articulation lines of urosome strong,
posteroventral setae of urosomite 1 much shorter
than in female. Spine formulas of uropods, uropod
1 peduncle apicolateral = 3, basofacial = 7-8, uro-
pod 2 peduncle dorsal = 9-12, dorsal spines on
outer ramus of uropod 1 = 3-4, of uropod 2 = 3?4,
inner ramus of uropod 1 = 1, of uropod 2 = 1
vestigial or 0. Ventral spines on peduncle of uropod
3 = 5; spine formula on article 1 of outer ramus
= 1-1-1-1-1-1, setal formula = 1-1-1-1-1-1. Tel-
son elongate, distal spines shortened.
OBSERVATIONS (male).?Male of Western Aus-
tralia, WAM 414-73, male "h": Right lacinia
mobilis free as in typical B. jirrandus; molar lack-
ing disjunct spine. Spine formula on article 4 of
antenna 2 = 1-4-3. Nail on dactyl of maxilliped 50
percent as long as article 4. Gnathopods slightly
stouter than in typical material (see illustrations of
outlines), poorly setose. Setal and spine formulas on
pereopods 1-2, article 4 = 3 and 2, article 5 = 4
and 4; article 6 = 3 + 3 plus middistal seta; article
5 of pereopod 1 lacking posteroproximal stout
spine, of pereopod 2 with one stout spine, dactyls
elongate, with faint inner acclivity bearing emer-
gent setule. Pereopods 3-5 with articles generally
thinner than in typical specimens (see illustrations
of outlines); anterior and posterior facial ridges on
pereopod 5 joined together, anterior ridge thus
elongate somewhat in resemblance of condition in
typical Birubius batei? (24). Epimeron 1 with 5-6
anteroventral setae, 2-3 posterofacial setae; epi-
meron 2 with 8 horizontal setae, none thrust dor-
sally; epimeron 3 with 6 posterior setae, 2 fully
ventral setal spines. Peduncle of uropod 1 with 4
basofacial setae, 3 apicolateral spines, outer ramus
with 2 dorsal spines, inner with one; peduncle of
uropod 2 with 10 spines, outer ramus with 2, inner
with one. Lateral margin of article 1 on outer
ramus of uropod 3 with spine formula of 1-1-2-2-2,
setae = 1-1-1-1-1.
This male has the typical right lacinia mobilis of
Birubius jirrandus but differs from that species in
the elongate anterior ridge on pereopod 5, the ab-
sence of a disjunct spine on the mandibular molar,
the elongate maxillipedal nail and the doubled
spines on the outer ramus of uropod 3. This male
merges to a large extent with Western Australian
males of B. batei?.
Male "m": Substantially like male "h" but right
mandibular molar widi small disjunct spine (left
348 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 173.?Birubius jirrandus, new species, holotype, female "a," 9.8 mm (m = male "m,
4.01 mm; v = male "v," 8.7 mm; a; = female "w," 103 mm; y = male "y," 8.1 mm).
NUMBER 245 349
aU
FIGURE 174.?Birubius jirrandus, new species, holotype, female "a," 9.8 mm (h = male "h," 4.00
mm; m ? male "m," 4.01 mm; w ? female "w," 10.5 mm).
350 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 175.?Birubius jirrandus, new species, female "f," 9.0 mm (to ? female "w," 103 mm;
y ? venter of urosomite 1).
NUMBER 245 351
side not examined), anterior ridge on pereopod 5
normal, disconnected from posterior ridge, right
epimeron 1 normal, left with 3 anteroventral setae
and only one posteroventral setule, posteroventral
tooth of epimeron 3 sharper than in other material
of this species (see illustration), telson left lobe
aberrant, perhaps regenerative (see illustration).
ILLUSTRATIONS.?Female left mandible drawn
from outwardly rolled position; inner plate of
maxilliped and article 3 of palp excessively flat-
tened; bases of missing setae on uropod 3 indicated
by circles; male inner plate of maxilla 2 drawn from
ventral view; most distal facial spine on article 5 of
pereopod 1 (Figure 175: wPl) obscured; male
pereopod 4 drawn in outline only for comparison
of proportions with female pereopod 4; setae on
uropod 3 actually thicker than shown in figure of
pleon; preservational defect in urosome marked
with arrow (occurs to a greater or less degree in
other males examined); female "w," 10.5 mm, exten-
sively illustrated to show condition in extremely
large individual, comparative reference to this speci-
men also made in text; mandibles, maxillae, and
maxilliped resembling those of holotype, not
illustrated; 2 posterior setae on epimeron 3 recon-
structed; dorsoposterior setule on epimera 1-2 occa-
sionally omitted in illustrations.
HOLOTYPE.?NMV, female "a," 9.8 mm .
TYPE-LOCALITY?CPBS 41S, 24 Mar 1965, West-
ern Port, Victoria, Australia, 16 m, coarse sand.
VOUCHER MATERIAL (all illustrated).?WAM
414-73: male "h," 4.00 mm; male "m," 4.01 mm.
PPBES 984/5: female "f," 9.0 mm. PPBES 984/4:
female "u," 9.0 mm. CPBS 42S/4: male "v," 8.7 mm.
CPBS 51S/1: female "w," 10.5 mm. PPBES 984/3:
male "y," 8.1 mm.
RELATIONSHIP.?Birubius jirrandus fits into Key
C of Birubius because of its simple right lacinia
mobilis but in other respects it lies outside the
species of that key and group because of the special
setation on epimeron 3. In that respect it resembles
but differs from B. batei? (24) in the simplicity of
the right lacinia mobilis, the short apical spines on
the telson of the female, the disjunct spine on the
molars, short anterior ridge on pereopod 5 and
short outer plate of the maxilliped, and, in the
majority of specimens, the lack of doubled spine
sets on the outer ramus of uropod 3.
Birubius jirrandus fits the description of Birubius
rostratus (Dana) presented by Stebbing (1906) and
Pirlot (1932) except that B. jirrandus has ventral
setae on epimeron 3 and only 9 spines on the outer
plate of maxilla 1 and has a somewhat more at-
tenuate rostrum than does B. rostratus. On the
other hand we can make no positive identification
because of our discovery in this study that ex-
tremely minute details such as right lacinia mobilis,
ridges on pereopods, disjunct spines on molars, and
numerous other items constitute the distinguishing
characters among phoxocephalids in the southwest-
ern Pacific (and probably other places) and these
details are unknown from earlier studies.
The right laciniae mobiles of B. batei? and B.
jirrandus demonstrate the stepwise loss of the prin-
cipal flabellate element in the complex; one must
suggest that B. jirrandus has lost the right lacinia
mobilis entirely and that the first raker spine has
also become reduced in size and now represents the
lacinia mobilis. Perhaps in B. batei? the right
lacinia mobilis is simply 2 fused raker spines with
complete loss of the true lacinia mobilis.
See "Remarks" under B. myallus {4).
MATERIAL.?CPBS, 12 samples from 11 stations
(15); WPBES, 9 samples from 6 stations (16);
PPBES, 13 samples from 5 stations (30); WAM, one
sample (2).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 1-19 m, sand, coarse sand. Western
Australia, Barrow Island, presumably neritic.
31. Birubius yorlunus, new species
FIGURES 176-178
DESCRIPTION OF FEMALE.?Head about 22 percent
of total body length, greatest width almost 60 per-
cent of length; rostrum constricted, narrow, elon-
gate, exceeding middle of article 2 on antenna 1.
Eyes medium to large, clear of pigment, ommatidia
especially large. Article 1 on peduncle of antenna 1
about 0.7 times as long as wide, about half as wide
as article 2, ventral margin with 5-6 setules, weakly
produced dorsal apex with 2 setules; article 2 about
0.55 times as long as article 1, with proximoventral
cycle of 5 setae; primary flagellum with 9-13 arti-
cles, about 0.7 times as long as peduncle, aesthetascs
poorly developed; accessory flagellum with 9-12
articles. Spine formula on article 4 of antenna 2 =
1-3-4-5 or 1-3-4-^ 1, dorsal margin with notch bear-
ing one seta and one spine, ventral margin with
352 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 176.?Birubius yorlunus, new species, holotype, female "a," 5.60 mm (b - male "b," 5.51
mm; c ? male "c," 5.50 mm).
NUMBER 245 S53
6 groups of 2-4 long to medium setae, one ventro-
distal long spine; article 5 about 0.72 times as
long as article 4, facial spine formula = 1-2, dorsal
margin naked, ventral margin with 2 sets of one
long seta, 2 ventrodistal long to medium spines, one
of these set subdistally; flagellum about 1.1-1.3
times as long as articles 4-5 of peduncle combined,
with 11-14 articles. Mandibles with strong palpar
hump; right incisor with 3 teeth; left incisor with
3 humps in 2 branches; right lacinia mobilis simple,
denticulate; left lacinia mobilis with 4 teeth, middle
teeth shortened; right rakers 9; left rakers 8-9;
molar in form of short protrusion demarcated
mainly by spines, right molar with 4 primarily long
spines, none strongly disjunct, left molar with 3-5
primarily long spines, none disjunct, each molar
with plume (not illustrated); palp article 1 short,
article 2 with one long inner apical seta and one
other short inner seta, article 3 over 0.9 times as
long as article 2, oblique apex with 9-11 spine-
setae, basofacial formula = 0-1 or 0-0. Inner plate
of maxilla 1 large, thin, bearing one long apical
pluseta, one shorter apicomedial seta, 2 apicolateral
slightly shorter setae; palp article 2 with 3 apical-
medial marginal spines and 4-5 lateral to medial
submarginal setae. Plates of maxilla 2 extending
equally, outer scarcely broader than inner, outer
with 2 apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with one large thick
apical spine, 2 apicofacial setae, 4 medial setae,
outer plate with 6 medial and apical spines, no
apicolateral setae; palp article 1 with one apicolat-
eral seta, article 2 with one apicolateral seta, medial
margin of article 2 moderately setose, article 3 with
4 facial setae, no lateral setae, nail of article 4 long,
with 2 accessory setules. Coxa 1 expanded distally,
anterior margin weakly concave; main ventral setae
of coxae 1-4 = 6-7-9-0, or 7-8-8-0 or 5-(6-7)-6-0
or 6-6-6-0 (or other variants), posteriormost seta of
coxae 1-3 as long as others; anterior and posterior
margins of coxa 4 strongly divergent, posterior
margin convex, posterodorsal corner rounded, pos-
terodorsal margin ordinary, concave, undulant,
width-length ratio of coxa 4 = 9:10. Long poster-
ior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = (5-6)-6-(5-6)-(4-6), short anter-
iors = 2-1-1-1, long anteriors = (2-4)-(3-5 4- 3
facial)-0-0, occasional straw-seta posteriorly on
pereopod 2. Gnathopods generally ordinary but
somewhat stout; width ratios of articles 5-6 on
gnathopods 1-2 = 26:41 and 26:41, length ratios
= 50:70 and 46:70; palmar humps medium,
palms strongly oblique; article 5 of gnathopods 1-2
ovate-triangular, posterior margin margin rounded,
almost lobate. Pereopods 1-2 similar, facial setae
formula on article 4 = 3-4 and 3, on article 5 = 5
and 5-6; main spine of article 5 extending to M. 75
on article 6, article 5 with l-(2) proximoposterior
spines; spine formula of article 6 = 6 + 8 and 6-7
+ 8 plus middistal seta, some spines especially long;
acclivity on inner margin of dactyls of pereopods
1-2 sharp, produced as tooth, emergent setule short,
midfacial pluseta short. Coxae 5-7 posteroventral
setule formula = 1-1-1. Articles 4-5 of pereopods
3-4 of ordinary width, facial spine rows dense,
facial ridge formula on article 2 of pereopods 3-5
= 0-2-2 (anterior short); width ratios of articles
2, 4, 5, 6, of pereopod 3 = 40:44:34:14, of pereo-
pod 4 = 60:38:24:10, of pereopod 5 = 80:19:18:8,
length ratios of pereopod 3 = 74:34:36:40, of
pereopod 4 = 82:45:35:52, of pereopod 5 =99:
23:23:27; article 2 of pereopod 5 not reaching
middle of article 5, ventral margin with 2 long
setae; medial apex of article 6 finely combed, bear-
ing 3 weak digital processes. Posteroventral corner
of epimeron 1 rounded, posterior margin straight,
anteroventral margin with 3-5 medium setae, pos-
teroventral face with 2 long setae set vertically;
posteroventral corner of epimeron 2 rounded, quad-
rate, weakly protuberant, posterior margin almost
straight, facial ridge very short, obsolescent, facial
setae = 7-8, posteriormost pair or triad set verti-
cally; posteroventral corner of epimeron 3 rounded
then with setule sinus, posterior margin straight,
with 2 setule notches, face with 2 oblique rows of
4 setae near ventral and posterior margin or as few
as one row of 5 plus one posteriorly and one rudi-
mentary; epimera 1?3 with small seta on postero-
dorsal margin set in weak notch. Urosomite 1 pro-
duced ventrally at base of uropod 1, articulation
line incomplete, short; urosomite 3 protuberant
dorsally. Rami of uropods 1-2 with articulate en-
larged apical nails, outer rami of uropods 1-2 and
inner ramus of uropod 1 with accessory nails, or
these poorly contiguous, outer ramus of uropod 1
with 5-6 dorsal spines, inner with 3, outer ramus of
uropod 2 with 4-6 dorsal spines, inner with 2 dorso-
medial spines; peduncle of uropod 1 with one api-
colateral spine and one basofacial seta, medially
with 4?5 marginal spines, apicalmost enlarged;
354 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Lu
FIGURE 177.?Birubius yorlunus, new species, holotype, female "a," 5.60 m (c ? male "c," 5.50
mm; g ? female "g," 4.S0 mm; k ? female "k," 4.57 mm).
NUMBER 245 S55
peduncle of uropod 2 with 5-6 dorsal spines, med-
ially with one small apical spine. Peduncle of uro-
pod 3 with 5 ventral spines, dorsally with one
lateral spine, one medial setule; rami feminine,
inner extending to M. 80 on article 1 of outer
ramus, apex with 1-3 setae, medial and lateral
margins naked, article 2 of outer ramus elongate,
0.30, bearing 2 short to medium setae, apicomedial
margin of article 1 naked, lateral margin with 4-5
acclivities, spine formula = l-2-2-2-2-(2), setal
formula = 0. Telson long, length-width ratio =
33:28, not fully cleft, each apex wide, rounded,
lateral acclivity deep, narrow, bearing ordinary
lateral setule, spine next medial of length equal to
setule, mid lateral setules diverse. Cuticle with ordi-
nary bulbar setules, surface bearing fine striations
in form of linear fingerprint pattern, emergent
setules especially short.
DESCRIPTION OF MALE.?Article 2 of antenna 1
with 5-6 ventral setae; primary flagellum 10-12
articulate, bearing calceolus on articles 1-4 or 1-5;
accessory flagellum 8-10 articulate. Facial spines on
article 4 of antenna 2 = 3-4-3 or 3-4-4, on article
5 = 2, ventrodistal apex of article 5 with 2 thin
spines, dorsal margin with 2-3 sets of male setae,
calceolus present or absent; flagellar formula =
(25-27 +), 2, 4, 6 . . . (20 or 26) (or other variants).
Right mandibular molar with 4 spines, left with 3;
left lacinia mobilis with 4 teeth plus one accessory
tooth; basofacial setal formula on article 3 = 1-1.
Inner plate of maxilliped with only 3 medial setae;
accessory setule on dactyl enlarged. Setal formula
on coxae 1-4 = 5-6-5-0; coxa 4 broader than in
female and with distinctive shape (see illustration).
Setal formulas on article 2 of gnathopods 1-2 and
pereopods 1-2, long posteriors = 5-5?4?4, long
anteriors = 2-3-0-0, short anteriors = 2?1?1?1-
Facial seta and spine formulas on pereopods 1-2,
article 4 = 2 and 2, article 5 = 3-4 and 4, article
6 = 6 + 8 plus middistal seta, posteroproximal on
article 5 = 1 and 1; posterodistal spine on article 5
of pereopods 1-2 slightly enlarged. Article 2 of
pereopods 3-4 narrower than in female. Setal
formulas on epimera, anteroventral epimeron 1 = 3 ,
epimeron 2 = 8, epimeron 3 = 6. Spine formulas on
uropods, apicolateral peduncle uropod 1 = 2 ,
peduncle of uropod 2 = 9, outer ramus = 4,
peduncle of uropod 3 = 8. Telson similar to that
of female but apical spines shorter, bearing dorsal
rows of denticles (mainly based on male "b").
OBSERVATIONS.?Routine identification of this
species demonstrates that the continuous dorsal
spination on the rami of uropods 1-2 is usually not
as strong as shown for the specimen depicted in our
illustrations. This is especially true of the inner
ramus on uropod I.
ILLUSTRATIONS.?Outer ramus of uropod 3 on
lateral view of pleon on male "c" added from inter-
polation of same ramus on male "b."
HOLOTYPE.?NMV, female "a," 5.60 mm.
TYPE-LOCALITY.?PPBES 974/2, 13 Oct 1971,
Port Phillip Bay, Victoria, Australia, 5 m, sand.
VOUCHER MATERIAL.?Type-locality: male "b,"
5.51 mm (illus.); male "c," 5.50 mm (illus.). CPBS
I2S/1: female "g," 4.30 mm (illus.). CPBS 12S/5:
female "k," 4.57 mm (illus.); male "m," 4.10 mm.
EBS 27: female "w," 5.7 mm.
RELATIONSHIP.?Birubius yorlunus combines the
following characters: simple right lacinia mobilis,
strong ventral setae on epimeron 1 with extra facial
setae in adults and poor to absent posterior setae,
absence of setae on coxa 4, numerous setae on
coxae 1-3, presence of accessory apical nails and
continuous spination on the rami of both uropods
1-2. In addition it has these minor attributes: un-
usual hands and palms of gnathopods, strong
proximal placement of setae on article 2 of antenna
1, small outer plate of the maxilliped, and short-
ened articles on the flagella of antenna 1. The lat-
ter characters can be utilized to draw comparisons
with the few other species bearing one or more of
those attributes. The following discussion is con-
fined to the major set of attributes in attempting to
establish relationships.
Birubius yorlunus falls between the B. myallus
(4) and B. taldeus (34) groups and bears comparison
to B. jirrandus (30). Birubius yorlunus differs from
the B. myallus group in the simple right lacinia
mobilis, the poorly developed basofacial setae on
the peduncle of uropod 1, the very thin rostrum,
and the short anterior ridge on article 2 of female
pereopod 5.
Birubius yorlunus shares with the B. taldeus
group the following characteristics: the poorly de-
veloped bulb on the anteroventral setules of coxa
\-?, similarities in right lacinia mobilis, thin
rostrum, and poorly developed basofacial setae on
uropod 1 but B. yorlunus is distinguished by the
strong ventral setae on epimeron 3, the multisetose
coxae 1-3, in the continuously spinose rami and
356 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
FIGURE 178.?Birubius yorlunus, new species, holotype, female "a," 5.60 mm (c ? male "c,1
5.50 mm; g ? male "g," 4.30 mm).
accessory nails of uropods 1-2, and in the distinctive
palms of the gnathopods. In all but uropods 1-2
B. yorlunus could serve as an hypothetical ancestor
to the B. taldeus group.
Birubius yorlunus differs from B. jirrandus in
the narrow rostrum, poor basofacial setae of uropod
1, absence of true posterior setae on epimeron 3,
and the presence of accessory apical nails and con-
tinuous dorsal spination on uropods 1-2.
The presence of accessory nails on uropod 2 sug-
gests that B. yorlunus is either more primitive than
any other member of Birubius in this one character
or that accessory nails can be developed late in the
evolutionary sequence.
Many specimens have the so-called "accessory"
apical nails, especially on the inner ramus of
uropod 1, less strongly developed and not as
crowded distally as in our illustrations, so that the
routine identificatory differences between B. yor-
lunus, B. eake (32), and B. kabbulinus (33) are often
NUMBER 245 357
obscured. Birubius eake can be distinguished from
B. yorlunus in the much shorter rostrum and much
more elongate wrists on the gnathopods, while B.
kabbulinus, known for the male only, can be dis-
tinguished in the absence of a distinct tooth and
sinus posteroventrally on epimeron 3 and in the
extreme basal position of the single dorsal spine on
the inner rami of uropods 1 and 2. These two char-
acters also distinguish B. kabbulinus from B. eake
as well as the longer rostrum and shorter wrists on
the gnathopods of B. eake.
MATERIAL.?CPBS, 6 samples from 2 stations
(252); PPBES, 32 samples from 18 stations (416);
EBS, one sample (1); SBS, 5 samples from 3 stations
(7)-
DISTRIBUTION.?Victoria, Western Port and Port
Phillip Bay to Sydney, New South Wales, 2-40 m,
sand and silty sand.
32. Birubius eake, new species
FIGURES 179-181
DESCRIPTION OF FEMALE.?Head about 19 percent
of total body length, greatest width about 67 per-
cent of length; rostrum constricted, narrow, short,
exceeding apex of article 1 on antenna 1. Eyes
medium, largely occluded with pigment. Article 1
on peduncle of antenna 1 about 1.15 times as long
as wide, twice as wide as article 2, ventral margin
with 6-7 setules, unproduced dorsal apex with one
setule; article 2 about 0.75 times as long as article 1,
with ventral cycle of 4 setae; primary flagellum with
8-9 articles, about 0.95 times as long as peduncle,
bearing long aesthetascs; accessory flagellum with
6 articles. Spine formula on article 4 of antenna
2 = 1-3-3-2, dorsal margin with notch bearing 2
setae, ventral margin with 3 groups of 2?3 long to
medium setae, one ventrodistal long spine; article
5 about 0.7 times as long as article 4, facial spine
formula = 1-2, dorsal margin naked, ventral
margin with 2 sets of 1-2 long to short setae, 2
ventrodistal long to medium spines; flagellum about
1.2 times as long as articles 4-5 of peduncle com-
bined, with 9 articles. Mandibles with strong palpar
hump; right incisor with 3 teeth; left incisor with
5 humps in 2 branches; right lacinia mobilis simple,
pointed, with marginal denticles; left lacinia
mobilis with 4 teeth plus one accessory tooth; right
rakers 4, plus one rudimentary; left rakers 5; molar
in form of elongate plaque, right molar with 6
primarily long spines plus plumose seta plus one
seta strongly disjunct, left molar with 3 primarily
long spines plus ciliate seta and one setal spine
strongly disjunct; palp article 1 slightly elongate,
article 2 with one long inner apical seta and one
other short inner seta, article 3 about 0.9 times as
long as article 2, oblique apex with 5 spine-setae,
basofacial formula = 0. Inner plate of maxilla 1
thin, bearing one long apical pluseta, one shorter
apicomedial seta, 2 apicolateral much shorter setae;
palp article 2 with 2 apicalmedial marginal spines
and 3 submarginal setae. Plates of maxilla 2 extend-
ing equally, outer slightly broader than inner, with
one apicolateral seta, inner with 2 medial setae.
Inner plate of maxilliped with one thin apical spine
and one partner seta, 2 apicofacial setae, 3 medial
setae; outer plate with 3 medial and apical spines,
no apicolateral setae; palp articles 1-2 with apico-
lateral seta, medial margin of article 2 weakly
setose, article 3 unprotuberant, with 2 facial setae,
2 lateral setae, nail of article 4 long, with 2 acces-
sory setules. Coxa 1 scarcely expanded distally,
anterior margin straight, main ventral setae of
coxae 1?4 = 4-4-4-0, posteriormost seta of coxae
1-3 as long as others; anterior and posterior
margins of coxa 4 weakly divergent, posterior
margin almost straight, posterodorsal corner
rounded, posterodorsal margin ordinary, concave or
V-shaped, width-length ratio of coxa 4 = 6:5. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 2-3-3-3, others not counted.
Gnathopods ordinary; width ratios of articles 5-6
on gnathopods 1-2 =27:37 and 27:37, length ra-
tios = 68:75 and 62:69; palmar humps ordinary,
palms oblique; article 5 of gnathopod 1 slightly
elongate, ovate, posterior margin rounded?flat; arti-
cle 5 of gnathopod 2 ovate, posterior margin
rounded, weakly lobate. Pereopods 1-2 similar,
facial setae formula on article 4 = 2 and 2, on arti-
cle 5 = 3 and 3; main spine of article 5 extending
to M. 80 on article 6, article 5 with one proximo-
posterior spine; spine formula of article 6 = 3 + 3
plus middistal seta; acclivity on inner margin of
dactyls of pereopods 1-2 obsolescent, emergent
setule long, midfacial pluseta ordinary. Coxae 5-7
posteroventral setule formula = 1-1-1. Articles 4?5
of pereopods 3-4 of ordinary width, facial spine
rows sparse, facial ridge formula on article 2 of
pereopods 3-5 = 0-2-1; width ratios of articles
S58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 179.?Birubius take, new species, holotype, female "a," 3.11 mm (x -? male "x," S.18 mm).
NUMBER 245 359
2, 4, 5, 6 of pereopod 3 = 46:41:34:16, of pereo-
pod 4 = 68:37:23:12, of pereopod 5 = 77:20:20:10,
length ratios of pereopod 3 = 73:27:33:41, of pe-
reopod 4 = 82:48:36:48, of pereopod 5 = 98:21:
21:29; article 2 of pereopod 5 slightly exceed-
ing apex of article 4; medial apex of article 6
finely combed and bearing 5 digital processes.
Posteroventral corner of epimeron 1 rounded, pos-
terior margin convex, with setule, anteroventral
margin with 3 short setae, posteroventral face with
one long seta; posteroventral corner of epimeron 2
rounded, posterior margin strongly convex, facial
setae = 7; posteroventral corner of epimeron 3
rounded, weakly and broadly protuberant, with
setule sinus, posterior margin weakly convex,
ventral face with horizontal crescent of 3 setae near
posterior margin. Urosomite I with long ventral
seta at base of uropod 1, articulation line short;
urosomite 3 unprotuberant dorsally. Rami of uro-
pods 1-2 with articulate enlarged apical nails, outer
ramus of uropod 1 with 2 dorsal spines, inner with
2, outer ramus of uropod 2 with 2 dorsal spines,
inner with one dorsomedial spine; peduncle of
uropod 1 with one apicolateral spine and one baso-
facial seta, medially with 2 marginal spines, apical-
most enlarged; peduncle of uropod 2 with 3 dorsal
spines, medially with one small apical seta. Pedun-
cle of uropod 3 with 4 ventral spines, dorsally with
one lateral spine, one medial spinule; rami fem-
inine, inner extending to M. 50 on article 1 of outer
ramus, apex with one seta, medial and lateral
margins naked, article 2 of outer ramus short, 0.15,
bearing 2 long setae, medial margin of article 1
with one seta, lateral margin with 2 acclivities,
spine formula = 2-2-2, setal formula = 0. Telson
ordinary, length-width ratio = 1:1, almost fully
cleft, each apex narrow, rounded, lateral acclivity
shallow, bearing ordinary lateral setule, spine next
medial longer than setule, midlateral setules di-
verse. Cuticle with ordinary bulbar setules.
DESCRIPTION OF MALE.?Eyes enlarged. Midface
of article 1 on antenna 1 bearing small patch of
fuzz; primary flagellum with 9 articles, one large
calceolus each on articles l^t; accessory flagellum
with 6 articles. Articles 3-4 of antenna 2 fuzzy
dorsally; spine formula of article 4 = 1-2-3-2, one
ventrodistal spine, ventral margin with 3 groups
of short to medium setae; article 5 with one
facial spine, 2 setules, 2 ventrodistal apical spines,
2 sets of short ventral setae, dorsal margin with one
calceolus and 2 sets of male setules; flagellar
formula = (21-22), 1, 2,4, 6 . . . 20. Right mandible
with 4 rakers plus one rudimentary; left with 4
rakers; right molar with 5 spines, one plumose
spine, one disjunct spine, left molar with 5 spines
plus one disjunct, no ciliated spine; right incisor
lacking fourth notch; mandibular palp with one
basofacial seta on article 3. Inner plate of maxilla
2 with one medial seta; outer plate with one apico-
lateral seta. Formula of long posteroventral setae on
coxae 1,2,3,4 = 3-3-3-0, coxa 4 bearing only one
posteroventral setule, otherwise coxa 4 like female.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = (l-2)-3-3-3. Pereopods 1-2
like female but article 5 of each with only 2 facial
setae. Article 2 of pereopod 3 relatively narrower
and longer than in female (see illustration), also
true for pereopod 4 but this appendage also larger
than in female; article 2 of pereopod 5 narrower
than in female, bearing 2 ridges. Epimera 1-3
broadened, posterior margin of epimeron 3 short-
ened; setal formulas, epimeron 1 anteroventral = 5,
posteroventral = 1, epimeron 2 facial = 7, epimeron
3 posterior = 2 setules, facial = 1, ventral = 1.
Ventral spine on each side of urosomite 1 much
shorter than in female and with peculiar falcate
shape; articulation line between urosomites 1-2
fully developed. Spine formulas of uropods, uropod
1 peduncle apicolateral = 2, basofacial = 1, uropod
2 peduncle dorsal = 6-7, dorsal spines on outer
ramus of uropod 1 = 2 , of uropod 2 = 2, inner
ramus of uropod 1 = 2, of uropod 2 = 1. Ventral
spines on peduncle of uropod 3 = 4, spine formula
on article 1 of outer ramus = 1-2-2-2, setal formula
= 1-1-1-1. Telson elongate, distal spines shortened.
Young male "n," 2.60 mm: Eyes scarcely larger
than those of female. Flagellum of antenna 2
slightly elongate, about 11-articulate, basal articles
short, spine formula on article 4 of peduncle =
1-3-3-2, article 5 with 2 facial spines. Epimeron 1
with 3 anteroventral setae, one posterofacial seta;
epimeron 2 with 6 facial setae; epimeron 3 with 3
facial setae. Peduncle of uropod 1 with one baso-
facial seta, 2 apicolateral spines, outer ramus with
2 dorsal spines, inner ramus with one dorsal spine;
uropod 2 with 3 peduncular spines, outer ramus
with 2 dorsal spines, inner ramus with one. Outer
ramus of uropod 3 with 2 acclivities, spine formula
= 2-2-2, setae absent on article 1.
DESCRIPTION OF JUVENILE (Juvenile "1," 2.00
360 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 180.?Birubius eake, new species, holotype, female "a," S.ll mm (x ? male "x," 3.18 mm).
NUMBER 245 361
FIGURE 181.?Birubius eake, new species, holotype, female "a," S.ll mm (X ? male "x," 3.18 mm).
mm).?Eyes small. Primary flagellum of antenna 1
with 7 articles; accessory flagellum with 5 articles.
Spine formula on article 4 of antenna 2 = 1-3-3-1;
article 5 with 2 facial spines. Epimeron 1 with one
anteroventral seta, no posterofacial seta; epimeron
2 with 4 facial setae; epimeron 3 with 2 facial setae.
Long setae of coxae 1-3 = 3-3-3. Ventral spine on
urosomite 1 thicker and larger relative to body size
than in adult female. Peduncle of uropod 1 with
one basofacial seta, one apicolateral spine, each
ramus with one dorsal spine; peduncle of uropod 2
with 2 spines, each ramus with one dorsal spine; all
spines of uropods 1-2 much larger than in adult
relative to body size, spines probably not increasing
steadily with body enlargement. Outer ramus of
uropod 3 with one lateral acclivity on article 1 bear-
ing one spine.
OBSERVATIONS.?Prebuccal complex convex an-
teriorly, moderately extended anteriorly; lower lip
with one cone on outer lobes, mandibular lobes
weakly subsharp and attenuate, like B. gambodeni
(20), epimeron 1 lacking lateral ridge in holotype,
probably obliterated by preecdysical swelling.
ILLUSTRATIONS.?Outer plate of maxilla 1 drawn
obliquely; male antenna 2 with one spine missing
on face of article 5.
362 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
HOLOTYPE.?WAM, female "a," 3.11 mm.
TYPE-LOCALITY.?JLB AUS 3, 1 Sep 1968, Sugar-
loaf Rock, Cape Naturaliste, Western Australia,
intertidal wash of common seaweeds.
VOUCHER MATERIAL.?Type-locality, male "x,"
3.18 mm (illus.); JLB AUS 12, small male "n," 2.60
mm; JLB AUS 13, juvenile "1," 2.00 mm.
RELATIONSHIP.?Birubius eake differs from the
other members of the B. taldeus (34) group in the
denser development of setae on coxa 1, the absence
of accessory apical setules on the rami of uropods
1-2, the presence of a few facial setae on epimeron
3, uncrowded setae on epimeron 2 possibly owing
to the narrower development of that plate, weaker
acclivity on the dactyls of pereopods 1-2, and nar-
rower articles 4-5 of either pereopod 3 or 4.
Birubius eake resembles B. munggai (27) but dif-
fers from that species in the simple right lacinia
mobilis, the noncrowded setae on epimeron 2 and
in the fewer setae of epimeron 3. From B. eleebanus
(29), B. eake differs in the fewer ventral setae on
epimeron 3, absence of setae on coxa 4 and the
fewer basofacial setae on uropod 1. See remarks on
this species under B. yorlunus (31).
We leave the reader the task of tracing the pos-
sible descent of this species from groups centered by
B. myallus (4), B. gallangus (10), B. narus (19), B.
gambodeni (20) and B. yorlunus (31). See other in-
terrelationships by tracing through from B. taldeus
(34), onward.
MATERIAL.?JLB AUS, 3 samples (7).
DISTRIBUTION.?Southwestern Australia, intertidal
flora.
33. Birubius kabbulinus, new species
FIGURES 182, 183
DESCRIPTION OF MALE.?Head (broken) about
21+ percent of total body length, greatest width
about 55 (estimated) percent of length; rostrum
constricted, narrow, elongate, probably reaching
middle of article 2 on antenna 1. Eyes large, clear
of pigment but stained deep burgundy, ommatidia
especially large. Article 1 on peduncle of antenna 1
about 1.2 times as long as wide, about twice as wide
as article 2, ventral margin with about 10 setules,
produced dorsal apex with one setule; article 2
about 0.8 times as long as article 1, with ventral
cycle of 4 setae; primary flagellum with 10 articles,
about 1.1 times as long as peduncle, lacking
aesthetascs; accessory flagellum with 7 articles. Spine
formula on article 4 of antenna 2 = 3-4-3, dorso-
medial margin of articles 3-4 fuzzy, ventral margin
with 4 groups of 1-3 medium setae, one ventrodistal
medium spine; article 5 about 0.9 times as long as
article 4, facial spine formula = 2, dorsal margin
bearing 3 sets of male setae and one calceolus,
ventral margin with 2 sets of 2 medium to short
setae, 2 thin vemrodistal medium spines; flagellum
elongate, flagellar formula = (26-29), 2, 4, 6 . . . 20
or 2, 4, 6 . . . 26. Mandibles with medium palpar
hump; right incisor with 3 teeth: left incisor with 4
humps in 2 branches; right lacinia mobilis simple,
denticulate; left lacinia mobilis with 5 teeth; right
rakers 5; left rakers 8; molar in form of bulbous
hump demarcated mainly by spines, each molar
with 4 primarily long spines, none disjunct, each
molar with plume; palp article 1 slightly elongate,
article 2 with one long inner apical seta and 2
other shorter inner setae, article 3 about 0.9 times
as long as article 2, oblique apex with 6 spine-setae,
basofacial formula = 1-1. Inner plate of maxilla 1
ordinary, bearing one long apical pluseta, one
shorter similar apicomedial seta, 2 apicolateral
much shorter setae; palp article 2 with 2 apical-
medial marginal spines and 3 submarginal setae.
Plates of maxilla 2 extending equally, of equal
width, outer with 2 apicolateral setae, inner with
one medial seta; [maxillipeds missing]; coxa 1
strongly expanded distally, anterior margin weakly
concave; main ventral setae of coxae 1-4 = 3-4-4-0,
posteriormost seta of coxae 1-3 longest; anterior
and posterior margins of coxa 4 almost parallel,
posterior margin convex, posterodorsal corner
rounded, posterodorsal margin short, concave, un-
dulant, width-length ratio of coxa 4 = 14:15. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 1-1-2-2, long anteriors = 2-5-0-0,
short anteriors = 1-1-1-1, no others. Gnathopods
ordinary; width ratios of articles 5-6 on gnathopods
1-2 = 24:36 and 26:37, length ratios = 56:60 and
49:59; palmar humps ordinary, palms oblique;
article 5 of gnathopod 1 ovate, posterior margin
rounded; article 5 of gnathopod 2 ovate, posterior
margin rounded, almost lobate. Pereopods 1-2 sim-
ilar; facial setae formula on article 4 = 2 and 2, on
article 5 = 3 and 2; main spine of article 5 extend-
ing to M. 85 on article 6, article 5 with one proxi-
moposterior spine; spine formula of article 6 = 5
NUMBER 245 S6S
FIGURE 182.?Birubius kabbulinus, new species, holotype, male "a," 3.19 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
R3
FIGURE 183.?Birubius kabbulinus, new species, holotype, male "a," 3.19 mm.
+ 3, lacking middistal seta, one spine especially
long; acclivity on inner margin of dactyls of pereo-
pods 1-2 weak, emergent setule long, midfacial
pluseta very short. Coxae 5-7 posteroventral setule
formula = 1?1-1. Articles 4-5 of pereopods 3-4 of
ordinary width, facial spine rows moderately devel-
oped, facial ridge formula on article 2 of pereopods
3-5 = 0-2-2 (see "Observations"); width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 39:37:30:12,
of pereopod 4 = 54:37:23:10, of pereopod 5 = 73:
17:15:7, length ratios of pereopod 3 = 60:26:
30:36, of pereopod 4 = 77:47:35:49, of pereopod
5 = 106:26:23:23; article 2 of pereopod 5 exceed-
ing apex of article 4; medial apex of article 6
combed, bearing 2 obsolescent digital processes.
Posteroventral corner of epimeron 1 rounded, pos-
terior margin deeply convex, anteroventral margin
with 4-5 medium setae, posteroventral face with 3
medium setae; posteroventral corner of epimeron
2 rounded? (damaged), posterior margin strongly
convex, facial setae = 6, posteriormost triad set
vertically; posteroventral corner of epimeron 3
rounded, with setule sinus, posterior margin convex,
with one setule notch, ventral margin with 5 setae
evenly spread; epimera 1-2 with tiny setule on
posterodorsal margin. Urosomite 1 naked, articula-
NUMBER 245 S65
tion line almost complete; urosomite 3 protuberant
dorsally. Rami of uropods 1-2 with articulate en-
larged apical nails, outer rami of uropods 1-2 and
inner ramus of uropod 1 with one accessory setule,
outer ramus of uropod 1 with 2 dorsal spines, inner
with one, outer ramus of uropod 2 with 3 dorsal
spines, inner with one dorsomedial spine; peduncle
of uropod 1 with 3 apicolateral spines and one
basofacial seta, medially with 5 marginal spines,
apicalmost weakly enlarged; peduncle of uropod 2
with 7 dorsal spines, medially with one apical
setule. Peduncle of uropod 3 with 5 ventral spines,
dorsally with one lateral spine, one medial setule;
rami masculine, inner extending to M. 100 on arti-
cle 1 of outer ramus, apex with 2-3 setae, medial
and lateral margins setose, article 2 of outer ramus
elongate, 0.25, bearing 2 long setae, apicomedial
margin of article 1 setose, lateral margin with 2
acclivities, spine formula = 1-1-1, setal formula =
1-1-1. Telson long, length-width ratio = 6:5, al-
most fully cleft, each apex narrow, rounded-
truncate, lateral acclivity weak, bearing ordinary
lateral setule, spine next medial shorter than setule,
midlateral setules weakly diverse. Cuticle with ordi-
nary but very sparse bulbar setules.
OBSERVATIONS.?Article 2 of pereopod 5 with 2
anterior ridges, one short and weak and possibly
an artifact; upper lip and lower lip poorly ob-
served, damaged during dissection; maxillipeds
missing.
HOLOTYPE.?AM, male "a," 3.19 mm. Unique.
TYPE-LOCALITY.?AM P. 18125, 12 Dec 1939,
Antechamber Bay, Kangaroo Island, South Aus-
tralia, light in net at night.
RELATIONSHIP.?Like B. eake (32), this species
lies just outside the B. taldeus (34) group. The right
lacinia mobilis is simple, the rostrum narrow, and
several rami of uropods 1-2 bear immersed apical
setules, but unlike the B. taldeus group, B. eake
and B. kabbulinus have more than 2 setae each on
coxae 1-2 and bear ventral or ventrofacial setal-
spines on epimeron 3. Birubus kabbulinus differs
from B. eake in the fully ventral position of the
setae on epimeron 3, bears an elongate article 2 on
the outer ramus of uropod 3, and lacks the enlarged
ventral spine on urosomite 1.
Birubius yorlunus (31) bears continuous dorsal
spination on the rami of uropods 1-2 and fully
facial setae on epimeron 3. Routine distinctions
however, are complicated; see B. yorlunus for re-
marks. Birubius kabbulinus also bears many simi-
larities to B. muldarpus (9) and B. gallangus (10)
but those species have bifid right lacinia mobili.
The absence of a distomedial seta on article 6 of
pereopods 1-2 is unusual.
Birubius kabbulinus has close resemblance to the
several taxa of the Cunmurra group, including
Birubius booleus (15) and B. chintoo (13). Birubius
kabbulinus differs from Cunmurra itickerus and
the two species 13 and 15 in the simple right lacinia
mobilis, from C. itickerus and B. chintoo in the
broad coxae 1-4, and in addition from Cunmurra
in the small gnathopod 2 and short hand of gna-
thopod 1. From B. booleus, B. kabbulinus also dif-
fers in the absence of long setae on coxa 4 and in
proportions of that coxa; however, the two species
are based on different sexes and other differences
may be masked by that factor.
Birubius kabbulinus does not fit the Cunmurra?
B. chintoo evolutionary scheme very well but its
mandibular molars have their spines reduced to 4
so that it closely approximates the genera of the
Paraphoxus-Brolgus group, though B. kabbulinus
does not have unusual gnathopods or combs on the
uropods.
MATERIAL.?AM, one sample (1).
DISTRIBUTION.?South Australia, Kangaroo Is-
land, neritic.
34. Birubius taldeus, new species
FIGURES 184-186
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about 65
percent of length; rostrum constricted, very narrow,
exceeding apex of article 1 on antenna 1. Eyes
medium, clear of pigment, stained ochre, omma-
tidia especially large. Article 1 on peduncle of an-
tenna 1 about 1.4 times as long as wide, about 1.6
times as wide as article 2, ventral margin with
8-10 setules, weakly produced dorsal apex with one
setule; article 2 about 0.7 times as long as article 1,
with ventral cycle of 4-5 setae; primary flagellum
with 12-13 articles, about 0.75 times as long as
peduncle, bearing short aesthetascs terminally; ac-
cessory flagellum with 8-10 articles. Spine formula
on article 4 of antenna 2 = 1-3-4-3 or 1-3-4--4,
dorsal margin with notch bearing one short spine,
ventral margin with 3 groups of 2 short to medium
366
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 184.?Birubius taldeus, new species, holotype, female "a," 3.5 mm (b -? male "b," 3.52
mm; m ? male "m," 3.7 mm).
NUMBER 245 367
FIGURE 185.?Birubius taldeus, new species, holotype, female "a," 3.5 mm (b = male "b," 332
mm; n = male "n," 2.83 mm).
setae, one ventrodistal long spine; article 5 about
0.8 times as long as article 4, facial spine for-
mula = 1-2, dorsal margin naked, ventral margin
with 2 sets of 2 medium to short setae, 2 ventro-
distal short to medium spines; ftagellum about 1.4
times as long as articles 4-5 of peduncle combined,
with 10-14 articles. Mandibles with medium to
strong palpar hump; right incisor with 3 teeth;
left incisor with 3 humps in 2 branches; right la-
cinia mobilis simple; left lacinia mobilis with 4
teeth; right rakers 7; left rakers 8; molar in form
of elongate plaque, right molar with 2-4 primarily
medium spines plus one very tiny disjunct spine,
left molar with 5 primarily medium spines plus
one tiny disjunct spine, each molar with plusetule;
palp article 1 slightly elongate, article 2 with one
medium inner apical seta and one other shorter
inner seta, article 3 about 0.75 times as long as
article 2, oblique apex with 5-6 spine-setae, baso-
facial formula = 0 (left), 1-1 (right). Inner plate
of maxilla 1 large, thin, bearing one long apical
pluseta, one shorter similar apicomedial seta, 2
apicolateral scarcely shorter setae, palp article 2
with 2 apicalmedial marginal spines and 3-4 sub-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 186.?Birubius taldeus, new species, holotype, female "a," 3.5 mm (b ? male "b," 352
mm; n ? male "n," 2.83 mm).
marginal setae, one of those apicolateral. Plates of
maxilla 2 extending equally, of equal width, outer
with 2 apicolateral setae, inner with one medial
seta. Inner plate of maxilliped with one large, thick
apical spine, 2 apicofacial setae, 3 medial setae;
outer plate with 5 medial and apical spines, no
apicolateral setae; palp article 1 lacking apicolat-
eral seta, article 2 with one apicolateral seta, me-
dial margin of article 2 weakly setose, article 3 pro-
tuberant, with 3 facial setae, no lateral setae, nail
of article 4 long, with 2 accessory setules. Coxa 1
strongly expanded distally, anterior margin almost
straight; main ventral setae of coxae 1-4 = 2-2-2-0,
posteriormost seta of coxae 1-3 shortened; anterior
and posterior margins of coxa 4 divergent, poste-
rior margin oblique, convex, posterodorsal corner
subsharp, posterodorsal margin short, concave, un-
dulant, width-length ratio of coxa 4 = 1:1. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 1-1-2-3, long anteriors = 1-1-0-0,
short anteriors = 2-3-3-3, no others. Gnathopods
ordinary; width ratios of articles 5-6 on gnatho-
pods 1-2 = 19:36 and 21:35, length ratios = 65:
56 and 58:53; palmar humps ordinary, palms
weakly oblique; article 5 of gnathopod 1 elongate,
narrowly ovate, posterior margin flat; article 5 of
gnathopod 2 ovate, posterior margin rounded-flat.
Pereopods 1-2 similar; facial setae formula on arti-
cle 4 = 1 and 1, on article 5 = 1 and 1; main spine
of article 5 extending to M. 75 on article 6, article
5 with one proximoposterior spine; spine formula
of article 6 = 3 + 4 plus middistal seta, one spine
especially long; acclivity on inner margin of dac-
tyls of pereopods 1-2 sharp, produced as tooth,
emergent setule long, midfacial pluseta ordinary.
Coxae 5-7 posteroventral setule formula = 1-1-1.
NUMBER 245 369
Articles 4-5 of pereopods 3-4 of ordinary width to
broad, facial spine rows moderately developed, fa-
cial ridge formula on article 2 of pereopods 5-5 =
0-2-2 (anterior short); width ratios of articles 2, 4,
5, 6 of pereopod 3 = 45:43:38:16, of pereopod
4 = 63:49:29:13, of pereopod 5 = 82:19:17:7, length
ratios of pereopod 3 = 76:30:37:40, of pereopod
4 = 87:53:41:50, of pereopod 5=117:26:22:29;
article 2 of pereopod 5 almost reaching apex of
article 5, ventral margin bearing medium sized
setae; medial apex of article 6 finely combed
and bearing 2 weak digital processes. Posteroven-
tral corner of epimeron 1 rounded-quadrate, pos-
terior margin convex, anteroventral margin with
one medium seta, posteroventral face with 2-3 me-
dium setae set vertically; posteroventral corner of
epimeron 2 rounded, posterior margin weakly con-
vex, facial setae = 4, weakly crowded, posterior-
most pair set vertically; posteroventral corner of
epimeron 3 rounded, then weakly protuberant,
with setule sinus and small tooth, posterior margin
almost straight, convex, serrate, setose (2-5 setae
besides posterodorsal seta), ventral margin naked;
epimera 1-3 with large seta on posterodorsal mar-
gin set in weak notch. Urosomite 1 naked, pro-
duced ventrally at base of uropod 1, articulation
line short; urosomite 3 unprotuberant dorsally.
Rami of uropods 1-2 with articulate enlarged api-
cal nails, all rami but inner ramus of uropod 2
with one accessory setule, outer ramus of uropod
1 with 4-5 dorsal spines, inner with 2-3, outer
ramus of uropod 2 with 2-3 dorsal spines, inner
with one dorsomedial spine; peduncle of uropod
1 with 2 apicolateral spines and one basofacial seta,
medially with 1-2 marginal setal spines plus apical
enlarged spine; peduncle of uropod 2 with 4-5 dor-
sal spines, medially with one small apical spine.
Peduncle of uropod 3 with 3-4 ventral spines, dor-
sally with 1-2 lateral spines, one medial setule;
rami feminine, inner extending to M. 60 on article
1 of outer ramus, apex with one seta, medial and
lateral margins naked, article 2 of outer ramus,
ordinary, 0.25, bearing 2 long setae, apicomedial
margin of article 1 naked, lateral margin with 3
acclivities, spine formula = 2-2-2-2, setal for-
mula = 0. Telson ordinary, length-width ratio =
12:11, not fully cleft, each apex of medium width,
rounded, lateral acclivity broad, shallow, bearing
ordinary lateral setule, spine next medial shorter
than setule, midlateral setules diverse. Cuticle with
ordinary bulbar setules emerging into clear halos
densely surrounded by short villi each bearing dark
spot or circle, villi especially prominent on body
segments, coxae 5-7, and on article 2 of pereopods
3-5, but absent or diminished on coxae 1-4, uro-
pods, and cephalad, emergent setules ordinary.
OBSERVATIONS (female).?Facial setae on article 3
of antenna 2 short. Female "o," 3.75 mm, PPBES
978/1, with 2 apicolateral spines on peduncle of
uropod 1.
DESCRIPTION OF MALE.?Eyes enlarged, dear of
occluding pigment, stained eosin in color. Article 2
of antenna 1 with 4 ventral setae; primary flagel-
lum with 9-12 articles, with one calceolus each on
articles 1-3 or 1-4, aesthetascs poorly developed;
accessory flagellum with 7-8 articles. Spine formula
on article 4 of antenna 2 = 3-4 4 or 1-2^4-3, ven-
tral setae of articles 4-5 short; distoventral corner
of article 5 with 1-2 short, thin spines plus or
minus setule, dorsal margin with 3 sets of male
setae, plus or minus one calceolus; flagellar for-
mula = (20-27), 2, 4, 6 . . . 22 (total 27) or 1, 2, 4,
6 . . . 20 (total 21) (or other variants). Right man-
dible with 6-7 rakers; left with 6-8; each molar
with 3-5 thin or flat spines and one stout spine,
no setule, plus extremely tiny disjunct spine; apical
seta on article 2 of palp elongate, right and left
palps with basofacial formula of 1-1 on article 3,
each apex with 6 spine-setae. Coxa 4 broader than
in female, posterodorsal margin longer. Setal for-
mula for article 2 of gnathopods 1-2 and pereopods
1-2, long posteriors = 2-2-3-3, long anteriors =
2-3-0-0 (plus one long facial on gnathopod 2),
short anteriors = 1-1-2-1, no others. Facial seta
formula on article 4 of pereopods 1-2 = 2 and 2,
on article 5 = 2 and 2-3, article 5 with one pos-
teroproximal spine. Pereopods 3-4 with narrower
articles than in female, or young male similar to
female, article 2 of pereopod 5 narrower, anterior
facial ridge weaker. Epimera 1-3 broadened, epi-
mera 1-2 extremely protuberant posteriorly, poste-
rior margin of epimeron 3 shortened; setal for-
mulas, epimeron 1 anteroventral = 2, posteroven-
tral = 2 vertical, epimeron 2 facial = 4, epimeron
3 posterior = 2-4 setules. Urosome slightly nar-
rower than in female, articulation line of urosomite
1 complete. Spine formulas of uropods, uropod 1
peduncle apicolateral = 1-2, basofacial = 1, uropod
2 peduncle dorsal = 7-8, dorsal spines on outer ra-
mus of uropod 1 = 3-5, of uropod 2 = 2-4, inner
370 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
raraus of uropod 1 = 1, of uropod 2 = 1 . Ventral
spines on peduncle of uropod 3 = 4-6, spine for-
mula on article 1 of outer ramus = 1-1-2-2-2, setal
formula = 1-1-1-1-1. Telson not elongate, distal
spines slightly shortened. Following parts like those
of female: maxillae, maxilliped, coxae 1-3.
OBSERVATIONS (male).?Male antenna 2 with
number of articles in flagellum variable and not
necessarily longer in terminal males than in
younger males.
ILLUSTRATIONS.?Following parts of female like
those illustrated for B. maldus (36) except in minor
details given in description: antenna 1, antenna 2,
prebuccal, upper lip (as shown for male of B. mal-
dus), maxillae 1-2, maxilliped, pereopodal dactyls;
mandibles generally like those of B. maldus with
differences illustrated for B. taldeus; male antenna
2 like male of B. maldus.
HOLOTYPE.?NMV, female "a," 3.5 mm.
TYPE-LOCALITY.?CPBS 61N/4, 23 Mar 1965,
Western Port, Victoria, Australia, 17 m, sand.
VOUCHER MATERIAL (all illustrated).?CPBS
42S/4, male "b," 3.52 mm; CPBS 61N/3, male "m,"
3.7 mm; CPBS 42S/4, male "n," 2.83 mm.
RELATIONSHIP.?The Birubius taldeus group con-
tains B. yandus (35), B. maldus (36) and B. eake
(32) and is characterized by: simple right lacinia
mobilis, very narrow rostrum, expanded coxa 1,
ordinary coxa 4 lacking long setae, and, in all but
B. eake, the reduction of the setae on coxae 1-3 to
2 in number, the absence of ventral or facial setae
on epimeron 3, and the presence of an accessory
apical setule on the rami of uropods 1-2. Epimeron
3 is straight to slightly convex posteriorly, has a
weak posteroventral protrusion with setule sinus,
and, except on B. eake, has 1-5 posterior setules.
The anteroventral setule on coxae 1-4 has the basal
bulb reduced in size or absent in the B. taldeus
group.
The B. taldeus group may have a point of origin
near B. yorlunus (31) a species bearing facial and
ventral setae on epimeron 3, fully developed acces-
sory nails on uropods 1-2 and fully developed setae
on coxa 1. However, this supposition would require
the hypothesis that the presence of apical setule-
scales on the rami of uropods 1-2 in the B. taldeus
group is a case of vestigial reduction in the arma-
ment, rather than the reverse hypothesis that ac-
cessory setules become accessory nails in an ordi-
nary pathway of evolution.
The B. taldeus group bears close resemblance to
B. munggai (27) but the right lacinia mobilis of
the B. taldeus group is simple, and except for B.
eake, epimeron 3 lacks facial setae and bears fewer
setae on epimeron 1.
Two of the species in the B. taldeus group
share with B. karobrani (14) the development of
ventral setae on article 2 of pereopod 5 but the
B. taldeus group otherwise differs from B. karo-
brani in the absence of setae on coxa 4, the simple
right lacinia mobilis, the poor basofacial setation
on uropod 1, and the absence of true ventral setae
on epimeron 3.
The B. taldeus group differs from B. eleebanus
(29) in the narrower and longer rostrum, the ab-
sence of setae on coxa 4, fewer setae on other ante-
rior coxae, no ventral setae on epimeron 3 (except
for B. eake) and the reduced basofacial setation on
uropod 1.
One may also trace ancestral lines for the B.
taldeus group through the following species or
groups: B. muldarpus-B. gallangus (9-10) and B.
mayamayi group (11-13) via epimeron 3; B. narus
(19) through coxa 1, the right lacinia mobilis and
the fully posterodorsal setule of epimeron- 3; B.
ularitus (28) in epimeron 3 and right lacinia mo-
bilis but not through coxa 1; and B. kabbulinus
(33) through epimeron 3 and coxa 1.
MATERIAL.?CPBS, 26 samples from 12 stations
(31); WPBES, 8 samples from 5 stations (11);
PPBES, 8 samples from 4 stations (24).
DISTRIBUTION.?Victoria: Western Port and Port
Philip Bay, 4-19 m, sand, silty sand.
35. Birubius yandus, new species
FIGURES 187-189
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total body length, greatest width about 60
percent of length; rostrum constricted, narrow,
elongate, almost reaching middle of article 2 on
antenna 1. Eyes medium to large, largely occluded
with pigment. Article 1 on peduncle of antenna 1
about 1.25 times as long as wide, about 1.7 times
as wide as article 2, ventral margin with about 6
setules, weakly produced dorsal apex with one set-
ule; article 2 about 0.7 times as long as article 1,
with proximoventral cycle of 4-5 setae; primary
flagellum with 10-11 articles, about 1.1 times as
NUMBER 845 371
FIGURE 187.?Birubius yandus, new species, holotype, female "w," 3.17 mm (/ ? female "f,"
3.17 mm; v ? female "v," 2.91 mm; y ?? male "y," 2-83 mm).
long as peduncle, bearing very few aesthetascs; ac-
cessory nagellum with 8-10 articles. Spine formula
on article 4 of antenna 2 = 1-3-4-3, dorsal margin
with notch bearing one seta and one spine, ventral
margin with 4 groups of 2 long to medium setae,
one ventrodistal long spine; article 5 about 0.75
times as long as article 4, facial spine formula =
1-2, dorsal margin naked, ventral margin with 2
sets of 1-2 long to short setae, 2 ventrodistal long
to medium spines; nagellum about 1.5 times as
long as articles 4-5 of peduncle combined, with
10-11 articles. Mandibles with strong palpar hump;
right incisor with 3 teeth; left incisor with 4 humps
in 2 branches; right lacinia mobilis simple; left
lacinia mobilis with 4-5 teeth, middle teeth weakly
shortened; right rakers 7-8, plus or minus one rudi-
mentary; left rakers 8; molar in form of elongate
plaque, right molar with 4 primarily long and one
rudimentary spines plus setule, none disjunct, left
molar with 5 primarily medium spines, none dis-
372 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Ek
FIGURE 188.?Birubius yandus, new species, holotype, female "w," 3.17 mm (/ ? female "f,1
3.17 mm; y ? male "y," 2.83 mm; N ? molar detail).
NUMBER 245 373
fW
FIGURE 189.?Birubius yandus, new species, holotype, female "w," 3.17 mm (/ ? female "f,1
S.I 7 mm; y ? male "y," 2.83 mm).
374 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
junct, each molar with plume; palp article 1 short,
article 2 with one long inner apical seta and one
other short inner seta, article 3 almost 0.9 times as
long as article 2, oblique apex with 6 spine-setae,
basofacial formula = 0. Inner plate of maxilla 1
ordinary, bearing one long apical pluseta, one
shorter apicomedial seta, 2 apicolateral scarcely
shorter setae; palp article 2 with 6 apical-lateral-
medial spines and setae. Plates of maxilla 2 extend-
ing equally, of equal width, outer with 2 apico-
lateral setae; inner with one medial seta. Inner
plate of maxilliped with one large thin apical
spine, 2 aoicofacial setae, 4 medial setae; outer
plate w :j i 5 medial and apical spines, no apico-
lateral setae; palp article 1 lacking apicolateral
seta, article 2 with one apicolateral seta, medial
margin of article 2 weakly setose, article 3 scarcely
protuberant, with 2-3 facial setae, no lateral setae,
nail of article 4 medium, with 1-2 accessory setules.
Coxa 1 strongly expanded distally, anterior margin
almost straight; main ventral setae of coxae 1-4 =
2-2-2-0, posteriormost seta of coxae 1-3 shortened;
anterior and posterior margins of coxa 4 divergent,
posterior margin oblique, almost straight, postero-
dorsal corner rounded, posterodorsal margin short,
weakly concave, width-length ratio of coxa 4 =
17:20. Long posterior setae on article 2 of gnatho-
pods 1-2 and pereopods 1-2 = 1-2-3-4, short an-
teriors = 2-2-2-2, long anteriors = 1-2-0-0, no
others. Gnathopods ordinary; width ratios of arti-
cles 5-6 on gnathopods 1-2= 18:35 and 21:38,
length ratios = 70:61 and 63:60; palmar humps
large, palms weakly oblique; article 5 of gnathopod
1 elongate, narrow, posterior margin flat, long;
article 5 of gnathopod 2 elongate, posterior margin
rounded-flat. Pereopods 1-2 similar; facial setae
formula on article 4 = 1 - 1 or 2-2, on article 5 =
3-3; main spine of article 5 extending to M. 80 on
article 6, article 5 with one proximoposterior spine;
spine formula of article 6 = 4 + 5 or 5 + 5 plus
middistal seta, one spine especially long; acclivity
on inner margin of dactyls of pereopods 1-2 sharp,
produced as tooth, emergent setule long, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule
formula = 2-1-1. Articles 4-5 of pereopods 3-4
broad, facial spine rows dense to moderately de-
veloped, facial ridge formula on article 2 of pereo-
pods 3-5 = 0-2-2 (anterior ridge on pereopod 5
short); width ratios of articles 2, 4, 5, 6 of pereopod
3 = 42:48:40:14, of pereopod 4 = 58:48:26:8, of
pereopod 5 = 85:19:18:7, length ratios of pe-
reopod 3 = 73:33:42:43, of pereopod 4 = 85:47:
34:45, of pereopod 5 = 1 1 1 2 3 2 3 : 2 8 ; article 2
of pereopod 5 exceeding middle of article 5,
ventral margin with long setae; medial apex of
article 6 finely combed, bearing 3 very weak digital
processes. Posteroventral corner of epimeron 1
rounded, posterior margin convex, anteroventral
margin with 1-2 medium setae, posteroventral face
with 2-3 medium setae set vertically; posteroventral
corner of epimeron 2 weakly protuberant, posterior
margin convex, facial setae = 4-6, posteriormost
pair set vertically; posteroventral corner of epi-
meron 3 broadly rounded, with setule sinus far
above, with small tooth, posterior margin weakly
to strongly convex, with 0-1 setule notch, ventral
margin naked; cpimera 1-3 with large seta on
posterodorsal margin set in weak notch. Urosomite
1 naked, produced ventrally at base of uropod 1,
articulation line incomplete; urosomite 3 protuber-
ant dorsally. Rami of uropods 1-2 with articulate
enlarged apical nails, outer rami of uropods 1-2
with accessory setule, outer ramus of uropod 1 with
3-5 dorsal spines, inner with 2-3, outer ramus of
uropod 2 with 3-4 dorsal spines, inner with 2
dorsomedial spines, outer rami continuously spi-
nose to apex; peduncle of uropod I with one apico-
lateral spine, and one basofacial seta, medially with
2 thin marginal spines plus apical enlarged spine;
peduncle of uropod 2 with 5-7 dorsal spines, me-
dially with pair of apical spines. Peduncle of uro-
pod 3 with 4-5 ventral spines, dorsally with one
lateral spine, one medial setule; rami feminine,
inner extending to M. 70 on article 1 of outer
ramus, apex with 1-2 setae, medial and lateral mar-
gins usually naked; article 2 of outer ramus elon-
gate, 0.33, bearing 2 medium to long setae, apico-
medial margin of article 1 lacking setae, lateral
margin with 2-3 acclivities, spine formula = 2-2-2
or 1-2-2-2, setal formula = 0. Telson ordinary,
length-width ratio =12:11, not fully cleft, each
apex of medium width, rounded, lateral acclivity
shallow, weak, bearing ordinary lateral setule, spine
next medial of length equal to setule, or much
longer in subadults, midlateral setules diverse. Cuti-
cle with ordinary bulbar setules surrounded by
sharp or blunt villi bearing central spots; emergent
setules ordinary.
OBSERVATIONS (female).?Female "f": Head mal-
formed, upper lip massively cornified (illustrated),
NUMBER 245 375
right molar with extra minute spines around short
blunt spine (illustrated), left molar with only 2
spines (illustrated), coxa 4 narrowed distally (illus-
trated), article 6 of pereopod 4 shortened (illus-
trated), urosome damaged, cuticle on coxae 5-7
and bases of pereopods 3-5 with villi reduced to
minute crescentic studs (illustrated); specimen prob-
ably most advanced female available and widely
illustrated but basic description including holotype
and other similar females.
DESCRIPTION OF MALE.?Rostrum slightly longer
and broader than in female. Primary flagellum of
antenna 1 with 11 articles, one calceolus each on
articles 2, 3, 4, aesthetascs poorly developed; acces-
sory flagellum 8-articulate. Formula on article 4 of
antenna 2 = 3-4-3, of article 5 = 2, ventral apex
of article 5 with 2 thin spines and setule, dorsal
margin with 3 sets of male setae; flagellar for-
mula = (24-25), 2, 4, 6 . . . 22 (total 24) or 3, 5,
7 . . . 21 (total 25). Article 3 of mandibular palp
with basofacial setal formula of 1-1; left lacinia
mobilis with 4 teeth. Palp of maxilla 1 with 5
setae and spines. Coxa 4 broader than in female,
smaller relative to size of coxa 1 than in female.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = 1-2-2-2, long anteriors = 1-
2-0-0, short anteriors = 2-3-2-1, no others. Pereo-
pods 1-2, article 4 with one facial seta, article 5
with 2, article 6 with 4 + 5 spines plus middistal
seta, article 5 with one proximoposterior spine each.
Article 2 of pereopods 3-5 narrower than in fe-
male. Epimera 1-3 broadened, epimera 1-2 strongly
protruding posteriorly, posterior margin of epi-
meron 3 shortened; setae shortened, setal formulas,
epimeron 1 anteroventral = 1, posteroventral = 2
vertical, epimeron 2 facial = 4, epimeron 3 poste-
rior ? 3. Urosome much smaller than in female.
Spine formulas of uropods, uropod 1 peduncle
apicolateral = 1, medial = 2, basofacial = 1, uropod
2 peduncle dorsal = 6, dorsal spines on outer ramus
of uropod 1 = 4, of uropod 2 = 4, inner ramus of
uropod 1 = 2, of uropod 2 = 3. Ventral spines on
peduncle of uropod 3 = 5, spine formula on article
1 of outer ramus = 2-1-2-2-2 or 2-2-2-2, setal for-
mula = 0-1-1-1-1 or 1-1-1-1. Telson slightly elon-
gate, distal spines scarcely shortened.
ILLUSTRATIONS.?Mandibles and maxillae of holo-
type not illustrated, mandibles of female "f" illus-
trated, maxillae generally like those of B. maldus
(36); setosity of gnathopods generally as in B. tal-
deus (34); part of urosome on female "i" omitted
owing to damage; attachment of inner ramus on
uropod 3 of female "f" reconstructed owing to
shriveling; calculations for pereopods 3-5 based on
holotype.
HOLOTYPE.?NMV, female "w," 3.17 mm.
TYPE-LOCALITY.?PPBES 960/2, 16 Feb 1971,
Port Phillip Bay, Victoria, Australia, 17 m, sand.
VOUCHER MATERIAL (all illustrated).?PPBES
960/3, female "f," 3.17 mm; PPBES 967/5, male
"y," 2.83 mm; PPBES 967/2, female "v," 2.91 mm.
RELATIONSHIP.?Within its subgroup of Birubius,
B. yandus is characterized by the evenly continuous
dorsal spination on the outer rami of uropods 1-2.
In the other species of the group a wide gap occurs
between the apex of the ramus and the first spine,
a gap wider than the distance between spines 1 and
2. Birubius yandus differs from B. maldus (36) in
the villose cuticle and from B. taldeus (34) in the
presence of only one apicolateral spine on the pe-
duncle of uropod 1 in adults.
MATERIAL.?PPBES, 14 samples from 4 stations
(59).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 5-17 m, sand.
36. Birubius maldus, new species
FIGURES 190-192
DESCRIPTION OF FEMALE.?Head about 20 per-
cent of total body length, greatest width about
55-60 percent of length; rostrum constricted, nar-
row, elongate, exceeding middle of article 2 on
antenna 1. Eyes medium to large, clear of pigment,
ommatidia especially large. Article 1 on peduncle
of antenna 1 about 1.3 times as long as wide, about
twice as wide as article 2, ventral margin with
about 9 setules, weakly produced dorsal apex with
one setule; article 2 about 0.7 times as long as arti-
cle 1, with ventral cycle of 4 setae; primary flagel-
lum with 10 articles, about 0.9 times as long as
peduncle, aesthetascs weakly developed; accessory
flagellum with 8-9 articles. Spine formula on arti-
cle 4 of antenna 2 = 1-3-4-3 (one hidden in illus-
tration), dorsal margin with notch bearing one seta
and one spine, ventral margin with 2 groups of
2-3 long to medium setae, one ventrodistal long
spine; article 5 about 0.7 times as long as article 4,
facial spine formula = 1-2, dorsal margin naked,
376 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
cS
FIGURE 190.?Birubius maldus, new species, holotype, female "a," 3.20 mm (6 ? female "b,1
3.21 mm; c = male "c," 3.10 mm).
NUMBER 245 377
bGlt
FIGURE 191.?Birubius maldus, new species, holotype, female "a," 3.20 mm (b ? female **b,'
3.21 mm; c = male "c," 3.10 mm).
ventral margin with 2 sets of 2-3 long to short
setae, 2 ventrodistal long to medium spines; flagel-
lum about 1.35 times as long as articles 4?5 of
peduncle combined, with 10 articles. Mandibles
with strong pal par hump; right incisor with 3
teeth; left incisor with 3 humps in 2 branches;
right lacinia mobilis simple, denticulate; left la-
cinia mobilis with 4 teeth; right rakers 7 plus one
rudimentary; left rakers 7-8; molar in form of
short protrusion or bulbous hump demarcated
mainly by spines, right molar with 4-5 primarily
long spines plus one short spine not disjunct, left
molar with 4 long spines, each molar with plume;
palp article 1 short, article 2 with one medium
inner apical seta and one other shorter inner seta,
article 3 about 0.9 times as long as article 2, oblique
apex with 6 spine-setae, basofacial formula = 0.
Inner plate of maxilla 1 thin, bearing one long
apical pluseta, one shorter apicomedial seta, 2
apicolateral scarcely shorter setae; palp article 2
with 2 apicalmedial marginal spines and 4 sub-
marginal setae. Plates of maxilla 2 extending
equally, of equal width, outer with 2 apicolateral
setae, inner with one medial seta. Inner plate of
maxilliped with one large, thick, apical spine, 2
apicofacial setae, 3 medial setae, outer plate with
4 medial and apical spines, no apicolateral setae;
palp article 1 lacking apicolateral seta, article 2
378 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 192.?Birubius maldtu, new species, holotype, female "a," 3.20 mm (b ? female "b,1
351 mm; c ? male "c," 3.10 mm; v - male "v," 3.27 mm).
NUMBER 245 379
with one apicolateral seta, medial margin of article 2
moderately to weakly setose, article 3 weakly pro-
tuberant, with 2 facial setae, no lateral setae, nail
of article 4 long, with 2 accessory setules. Coxa 1
expanded distally, anterior margin straight; main
ventral setae of coxae 1-4 = 2-2-2-0, posteriormost
seta of coxae 1-3 slightly shortened; anterior and
posterior margins of coxa 4 almost parallel, poste-
rior margin very convex, posterodorsal corner
rounded, posterodorsal margin short, concave,
width-length ratio of coxa 4 = 1:1; long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 2-3-3-3, long anteriors = 2-3-0-0, short an-
teriors = 1-1-3-1, no others. Gnathopods ordinary;
with ratios of articles 5-6 on gnathopods 1-2 =
22:36 and 25:36, length ratios = 65:60 and 60:61;
palmar humps large, palms oblique; article 5
of gnathopod 1 elongate, ovate, posterior margin
rounded-flat, article 5 of gnathopod 2 ovate, pos-
terior margin rounded. Pereopods 1-2 similar; fa-
cial setae formula on article 4 = 2 and 2, on article
5 = 2 and 2, or 3 and 3; main spine of article 5
extending to M. 80 on article 6, article 5 with one
proximoposterior spine; spine formula of article
6 = 3 + 4 and 4 + 4 plus middistal seta, one spine
especially long; acclivity on inner margin of dac-
tyls of pereopods 1-2 sharp, produced as tooth,
emergent setule long, midfacial pluseta short.
Coxae 5-7 posteroventral setule formula = 1?1-1.
Articles 4-5 of pereopods $-4 of ordinary width to
broad, facial spine rows moderately developed, fa-
cial ridge formula on article 2 of pereopods 3-5 =
0-2-2 (anterior short on pereopod 5); width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 45:46:39:15,
of pereopod 4 = 60:47:28:11, of pereopod 5 =
80:19:15:7, length ratios of pereopod 3 = 75:
31:37:41, of pereopod 4 = 84:47:32:45, of pereopod
5 = 115:25:21:28; article 2 of pereopod 5 reach-
ing middle of article 5; medial apex of arti-
cle 6 finely combed and bearing 2 obsolescent digi-
tal processes. Posteroventral corner of epimeron 1
rounded-quadrate, posterior margin straight, ante-
roventral margin with one medium seta, postero-
ventral face with 2 long setae set vertically; postero-
ventral corner of epimeron 2 rounded-quadrate,
posterior margin straight, facial setae = 4-5, pos-
teriormost pair set vertically; posteroventral corner
of epimeron 3 rounded, undulant, weakly protuber-
ant, with setule sinus, posterior margin almost
straight, ventral margin naked; epimera 1-3 with
setule on posterodorsal margin. Urosomite 1 pro-
duced ventrally at base of uropod 1, articulation
line short; urosomite 3 unprotuberant dorsally.
Rami of uropods 1-2 widi articulate enlarged api-
cal nails, outer rami of uropods 1-2 with one ac-
cessory setule, outer ramus of uropod 1 with 3
dorsal spines, inner with 2, outer ramus of uropod
2 with 3 dorsal spines, inner with 2 dorsomedial
spines; peduncle of uropod 1 with one apicolateral
spine and one basofacial seta, medially with one
marginal seta and apical enlarged spine; peduncle
of uropod 2 with 4 dorsal spines, medially with one
small apical spine. Peduncle of uropod 3 with 3
ventral spines, dorsally with one lateral spine, one
medial setule; rami feminine to submasculine, inner
extending to M. 75 on article 1 of outer ramus,
apex with 1-2 setae, medial and lateral margins
naked, article 2 of outer ramus ordinary, 0.22, bear-
ing 2 medium to long setae, apicomedial margin of
article 1 naked, lateral margin with 3 acclivities,
spine formula = 1-2-2-2, setal formula = 0. Tel-
son ordinary, length-width ratio = 12:11, not fully
cleft, each apex of medium width, rounded, lateral
acclivity narrow, weak, bearing long lateral setule,
spine next medial of length equal to setule, mid-
lateral setules diverse. Cuticle with ordinary bulbar
setules, setules especially short.
DESCRIPTION OF MALE.?Article 2 of antenna 1
with 4?6 ventral setae; primary flagellum with 11
articles, one calceolus each on articles 1-5 or 1-6,
aesthetascs very poorly developed; accessory flagel-
lum with 9 articles. Facial formula on article 4 of
antenna 2 = 3-4-3, on article 5 = 2, article 5 with
3 dorsal sets of male setae and 0-1 calceolus, disto-
ventral corner with 1-2 thin spines and 0-1 setule;
flagellar formula, (25-26), 1, 2, 4, 6 . . . (25 or 26),
or 2, 4, 6 . . . . or 2, 3, 5, 7 . . . . Apical seta on
article 2 of mandibular palp more elongate than
in female, basofacial setal formula on article 3 =
1-1; right rakers = 8 plus one rudimentary; left
rakers = 9; each molar with 5 spines and plume.
Inner plate of maxilla 1 with only one apicolateral
seta instead of 2 as in female; palp with only 5
spines and setae. Outer plate of maxilliped with 6
spines (2 forming pair). Coxa 4 relatively broader
than in female, coxa 1 larger in relation to coxa 4
than in female. Long posterior setae on article 2
of gnathopods 1-2 and pereopods 1-2 = 1-1-2-3,
long anteriors = 1-2-0-0, short anteriors = 2-2-2-2,
no others. Facial setae on article 4 of pereopods
380 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
1-2 = 1 and 1-2, on article 5 = 2 and 2, spines on
article 6 = 3 + 4 and 4 + 4 plus middistal seta,
article 5 with one posterior spine on both pairs.
Article 2 of pereopods 3-5 narrower than in fe-
male, articles 4?5 of pereopod 4 more elongate,
width ratios of articles 5-6 = 26:11, length ra-
tios = 40:54 (in same enlargement as female). Epi-
mera 1-3 broadened, posterior margin of epimera
1-2 strongly protuberant, posterior margin of epi-
meron 3 shortened; setal formulas, epimeron 1 an-
teroventral = 1-2, posteroventral = 2 horizontal or
vertical, epimeron 2 facial = 3-5, epimeron 3 poste-
rior = 2. Urosome smaller than in female, articula-
tion line of urosomite 1 almost complete. Spine
formulas of uropods, uropod 1 peduncle apicolat-
eral = 1-2, basofacial = 1, uropod 2 peduncle dor-
sal = 8 (12 in supermale), dorsal spines on outer
ramus of uropod 1 = 4, of uropod 2 = 3-4, inner
ramus of uropod 1 = 2, of uropod 2 = 1 . Ventral
spines on peduncle of uropod 3 = 4 (7 in super-
male), spine formula on article 1 of outer ramus =
1-1-2-2-2, or 1-2-2-2-2, setal formula = 1-1-1-
1-1. Telson elongate, distal spines shortened. Bul-
bar setules on cuticle much sparser than in female
but cuticle becoming covered with fingerprint stria-
tions, grosser in terminal males.
OBSERVATIONS.?Larger or older males and fe-
males with apices on outer rami of uropods 1-2
becoming attenuate, distance between apex and
first distal spine becoming greater and greater; fe-
males "a" and "b" each with one egg in brood
pouch, female "a" with additional object, perhaps
rotted egg.
ILLUSTRATIONS.?Female upper lip shown with-
out epistomal part; male upper lip drawn with
epistomal part, entire figure drawn from slightly
oblique dorsal view, but male without ventral
hump; illustration of right (to viewer) side of lower
lip with outer margin folded.
HOLOTYPE.?NMV, female "a," 3.20 mm.
TYPE-LOCALITY.?CPBS 12S/3, 18 Mar 1965,
Western Port, Victoria, Australia, 2 m, muddy
sand.
VOUCHER MATERIAL (all illustrated).?Type-local-
ity, female "b," 3.21 mm (with much enlarged eyes);
CPBS 51N/1171, male "c," 3.10 mm; PPBES 974/1,
supermale "v," 3.27 mm.
RELATIONSHIP.?This species differs from B. tal-
deus (34) and B. yandus (35) in the nonvillose cuti-
cle and the poor ventral setation on article 2 of
pereopod 5. From B. yandus, in addition, it differs
in the wide gap between the apex and first distal
spine on the outer rami of uropods 1-2. From B.
taldeus, in addition, it differs in the parallel ante-
rior and posterior margins of coxa 4 in the female,
the presence of only one apicolateral spine on the
peduncle of uropod 1 (but supermale B. maldus
bears 2 spines and females of B. taldeus often have
only 1 spine), the longer spines on the telson,
presence of 2 elements in dorsal notch on article 4
of female antenna 2, elongate apical setae on the
inner ramus of uropod 3 of the female, narrower
coxae 1-2, longer spines on the peduncle of uropod
2 and the longer rostra and larger eyes in both
sexes.
MATERIAL.?CPBS, 13 samples from 7 stations
(22); WPBES. 2 samples from 2 stations (2); PPBES,
19 samples from 10 stations (30).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 2-25 in, sand, muddy sand, clayey silt,
clay.
37. Birubius wulgaru, new species
FIGURES 195-198
DESCRIPTION OF FEMALE.?Head about 15 per-
cent of total body length, greatest width 65-75
percent of length; rostrum obsolescent, scarcely
reaching along article 1 on antenna 1. Eyes me-
dium-large, largely occluded with pigment, omma-
tidia especially large. Article 1 on peduncle of an-
tenna 1 about 1.4 times as long as wide, about 1.75
times as wide as article 2, ventral margin with
about 5-6 setules, weakly produced dorsal apex
with 1-3 setules; article 2 about 0.75 times as long
as article 1, with ventral cycle of 6 setae; primary
flagellum with 10-11 articles, about 1.2-1.3 times
as long as peduncle, lacking aesthetascs; accessory
flagellum with 8 articles. Spine formula on article
4 of antenna 2 = 1-3-4 -4, dorsal margin with notch
bearing one spine, ventral margin with 2-3 groups
of 2-4 long or medium setae, one ventrodistal long
spine; article 5 about 0.7 times as long as article 4,
facial spine formula = 1-2, dorsal margin naked,
ventral margin with 2 sets of 1-2 long to short
setae, 2 other ventrodistal long to medium spines
disjunct or not from apicoventral spine; flagellum
1.5-1.8 times as long as articles 4-5 of peduncle
combined, with 11 articles. Mandibles with me-
NUMBER 245 381
FIGURE 193.?Birubius wulgaru, new species, female "c," 3.3 mm (z ? male "z," 3.4 mm).
382
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 194.?Birubius wulgaru, new species, female "c," 3.3 mm (z = male "z," 3.4 mm).
dium palpar hump; right incisor with S teeth; left
incisor with 3-4 humps in 2 branches; right lacinia
mobilis simple, denticulate; left lacinia mobilis
with 4 teeth plus one accessory tooth, middle teeth
shortened; right rakers 6; left rakers 7; molar in
form of elongate plaque, weakly bulbous, each
molar with 6 primarily long spines plus one tiny
spine strongly disjunct, usually each molar with
plume; palp article 1 slightly elongate, article 2
with one long to short inner apical seta and one
other shorter inner seta, article 3 about 0.7-0.8
times as long as article 2, oblique apex with 6
spine-setae, basofacial formula = 0. Inner plate of
maxilla 1 ordinary to large, bearing one long api-
cal pluseta, one shorter similar apicomedial seta,
2 apicolateral shorter setae, palp article 2 with 5
apicalmedial-lateral marginal spines and setae.
Plates of maxilla 2 extending subequally, outer
NUMBER 245 383
W
FIGURE 195.?Birubius wulgaru, new species, female "c," 3.3 mm (z =- male "z," 3.4 mm).
384 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 196.?Birubius wulgaru, new species, holotype, female "a," 3.42 mm (b ? male "b,1
3.49 mm).
NUMBER 245 385
FIGURE 197.?Birubius wulgaru, new species, holotype, female "a," 3.42 mm (b ?= male "b,"
3.49 mm).
386 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 198.?Birubius wvlgaru, new species, holotype, female "a," 3.42 mm (b ? male, "b.
3.49 mm).
NUMBER 245 387
narrower than inner, outer with 2 apicolateral
setae, inner with one medial seta. Inner plate of
maxilliped with one large, thick apical spine, 2-3
apicofacial setae, 4 medial setae in L-pattern; outer
plate with 2-5 medial and apical spines, 1-2 apico-
lateral setal spines, and weak to strong apical
spout; palp article 1 lacking apicolateral seta, arti-
cle 2 with one apicolateral seta, medial margin of
article 2 moderately setose, article 3 protuberant,
with 3 facial setae, no lateral setae, nail of article
4 long, with 2 accessory setules. Coxa 1 strongly
expanded distally, anterior margin straight; main
ventral setae of coxae 1-4 = 2-2-2-0, posteriormost
seta of coxae 1-3 shorter; anterior and posterior
margins of coxa 4 divergent, posterior margin very
oblique, convex, posterodorsal corner rounded,
posterodorsal margin long to ordinary, straight,
width-length ratios of coxa 4 = 7:8 or 9:10. Long
posterior setae of gnathopods 1-2 and pereopods
1-2 = l-(l-2)-2-3, short anteriors = 2-2-4-4, long
anteriors = 1-2-0-0, no others. Gnathopods with
elongate wrists and broadened hands; width ratios
of articles 5-6 on gnathopods 1-2 = 20:38 and
19:39, length ratios = 73:65 and 65:60; palmar
humps large, palms oblique; article 5 of gnathopod
1 ovate, posterior margin flat, long; article 5 of
gnathopod 2 ovate, posterior margin rounded-flat.
Pereopods 1-2 similar; facial setae formula on arti-
cle 4 = 1 and 1, on article 5 = 1 and 1; main spine
of article 5 extending to M. 80-95 on article 6,
article 5 with one proximoposterior spine; spine
formula o? article 6 = 3 + 4 and/or 4 + 4 plus
middistal setule, one spine especially long; acclivity
on inner margin of dactyls of pereopods 1-2 sharp,
produced as tooth, emergent setule long, midfacial
pluseta ordinary. Coxae 5-7 posteroventral setule
formula = 1-1-1. Articles 4?5 of pereopods 3-4
broad, facial spine rows dense to moderately de-
veloped, facial ridge formula on article 2 of pereo-
pods 3-5 = 0-2-2, anterior ridge on pereopod 5
short; width ratios of articles 2, 4, 5, 6 of pereopod
3 = 42:42:34:14, of pereopod 4 = 55:42:26:13, of
pereopod 5 = 78:17:16:16, length ratios of pereo-
pod 3 = 73:30:35:43, of pereopod 4 = 78:45:37:45,
of pereopod 5 = 106:27:24:30; article 2 of pereopod
5 reaching apex or middle of article 5, ventral
margin of article 2 moderately setose; medial
apex of article 6 finely combed and bearing
obsolescent digital processes. Posteroventral corner
of epimeron 1 rounded-quadrate or weakly
protuberant, posterior margin straight to weakly
convex, anteroventral margin with 2 long setae,
posteroventral face with 0-1 long seta; posteroven-
tral corner of epimeron 2 weakly protuberant, with
small, sharp tooth, posterior margin convex, facial
setae = 3-4, posteriormost pair set vertically; pos-
teroventral corner of epimeron 3 rounded, then
weakly protuberant, with setule sinus, posterior
margin weakly convex, with 0-1 setule notch,
ventral margin naked; epimera 1-3 with large seta
on posterodorsal margin set in weak to deep notch,
epimeron 3 thus with 2-3 posterior setae and
setules. Urosomite 1 naked, articulation line ob-
solescent or incomplete, often shifted anteroven-
trally; urosomite 3 unprotuberant dorsally. Rami
of uropods 1-2 with articulate enlarged apical nails,
outer rami of uropods 1?2 and inner ramus of
uropod 1 with one accessory setule, outer ramus of
uropod 1 with 4?5 dorsal spines, inner with 2-3,
outer ramus of uropod 2 with 3 dorsal spines, inner
with 1-2 dorsomedial spines; peduncle of uropod 1
with 1-2 apicolateral spines and one basofacial seta,
medially with 2-3 marginal setae and spines, apical-
most an enlarged spine; peduncle of uropod 2 with
4?5 dorsal spines, medially with one small apical
spine. Peduncle of uropod 3 with 3-4 ventral spines
and occasional extra proximal spine, dorsally with
one lateral spine, one medial setule; rami feminine,
inner extending to M. 70 on article 1 of outer
ramus, apex with one seta, medial and lateral mar-
gins naked, article 2 of outer ramus elongate, 0.30,
bearing 2 long setae, apicomedial margin of article
1 with 0-1 seta, lateral margin with 3-5 acclivities,
spine formula = 2-2-2-2 or 1-2-2-2-2-2, setal
formula = 0. Telson ordinary, length-width ratio
= 12:11, not fully cleft, each apex of medium
width, rounded-truncate, lateral acclivity deep,
broad, bearing long lateral setule, spine next medial
longer than setule, and second shorter spine next
medial, midlateral setules diverse. Cuticle with ordi-
nary bulbar setules surrounded by clear halo in
midst of weak to dense villose-scale pebbling,
emergent setules ordinary.
OBSERVATIONS (female).?Female "c": Width ra-
tios of articles 5-6 of gnathopods 1-2 = 22:39 and
22:40, length ratios = 75:65 and 68:61. Posterior
margin of article 4 on pereopod 1 with 7 setae be-
tween M. 50 and M. 100, on pereopod 2 with 9
setae between M. 33 and M. 100. Width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 53:55:46=17, of
388 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pereopod 4 = 68:58:32:17, of pereopod 5 = 93:
22:18:9, length ratios of pereopod 3 = 83:27:40:
47, of pereopod 4 = 95:60:43:50, of pereopod 5 =
120:30:26:35. Articulation line of urosomite 1
almost absent. Lateral margin on article 1 of
uropod 3 with 3 acclivities, spine formula = 2-
2-2-2. Lateral acclivity on telson weak, occasionally
one lobe with third small spine.
DESCRIPTION OF MALE.?Rostrum more attenuate
than in female. Article 2 of antenna 1 with 8
ventral setae; flagella elongate; primary flagellum
with 12-13 articles, one calceolus each on articles
1-4, aesthetascs absent; accessory flagellum with 8
articles. Facial spine formula on article 4 of antenna
2 = 1-2-4-4, on article 5 = 2; article 5 with 3
dorsal sets of male setae, no calceoli, ventrodistal
apex with 2 thin spines and one sctule; flagcllar
formula = 25, 2, 4, 6 . . . 22. Right rakers 7 plus one
rudimentary; left rakers 8 plus one rudimentary;
left lacinia mobilis with middle teeth obsolescent;
right molar with 6-8 spines plus weak plume,
usually no disjunct spine, left molar with 5-6
spines, weak plume and disjunct spine or not, baso-
facial setal formula on article 3 of mandibular palp
= 1-1, terminal = 6. Coxa 4 broadened but smaller
in relation to coxa 1 than in female. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = l-2-2-(2-3), long anteriors = 0-2-0-0, short
anteriors = 2-2-4-3, no others. Spine formula of
pereopods 1-2 on article 6 = 4 + 4 plus middistal
seta, on posteroproximal margin of article 5 = 1?2.
Article 2 of pereopods 3-5 not distinctly narrower
than in female but posterior margin of pereopod 5
flattened. Epimera 1-3 broadened, posterior margin
of epimeron 3 shortened; setal formulas, epimeron
1 anteroventral = 1, posteroventral = 1, epimeron
2 facial = 3, epimeron 3 posterior = 2-3 setules,
dorsoposterior setule on epimera 1-3 small. Spine
formulas of uropods, uropod 1 peduncle apicolat-
eral = 2, basofacial = 1, uropod 2 peduncle dorsal
= 8, dorsal spines on outer ramus of uropod 1 = 4 ,
of uropod 2 = 4 , inner ramus of uropod 1 = 2, of
uropod 2 = 1-2. Ventral spines on peduncle of uro-
pod 3 = 4, spine formula on article 1 of outer
ramus = 1-2-2-2-2-2 or 1-1-2-2-2-2, setal formula
= 1-1-1-1-1-1. Telson elongate, distal spines
slightly shortened.
OBSERVATIONS.?Cuticular villi densest on pereo-
pods 3-5 and dorsally on pleonites 1-3, sparse on
epimera, decreasing in density anteriorly on body
segments and absent on anterior body segments, ab-
sent on urosome except dorsally: note illustrations
of truncate spines on pereopods (3^4); each descrip-
tion based on 2 females and 2 males, numerous
minor details distinctive as shown in illustrations.
ILLUSTRATIONS.?Holotype, female "a": left molar
with dotted setule present only on right molar;
apex of palp on maxilla 1 not fully flattened. Fe-
male "c": several pereopods illustrated from right
side because of damage on left side of specimen.
Male "z": left molar enlargement showing disjunct
spine but only one spine of elongate group shown
for simplicity.
HOLOTYPE.?NMV, female "a," 3.42 mm.
TYPE-LOCALITY? WPBES 1704/2, 22 Nov 1975,
Western Port. Victoria, Australia, 12 m, clayey sand.
VOUCHER MATERIAL (all illustrated).?Type-local-
ity: male "b," 3.49 mm. PPBES 980 3: female "c,"
3.3 mm; male "z," 3.1 mm.
RELATIONSHIP.?Birubius wulgaru is an extension
of the B. taldeus (34) group. Strong resemblance oc-
curs in the right lacinia mobilis, coxae 1-4, cuticle,
epimera 1-3, and basofacial setae of uropod 1.
Birubius wulgaru differs from the B. taldeus group
in the much shorter rostrum and presence of two
spines on each apex of the telson (in contrast to one
spine on each apex).
Birubius wulgaru might be more directly
descendent from B. eleebanus (29) than from the
B. taldeus group. There is a strong resemblance be-
tween the two species in cephalic reduction, shapes
of gnathopods, cuticle, maxilla 2, and the posterior
margin of epimeron 3 but B. wulgaru differs from
B. eleebanus in the strong reduction of coxal setae,
much stouter lateral spines on article 1 of the outer
ramus on uropod 3 and the absence of ventral setae
on epimeron 3.
MATERIAL.?WPBES, one sample (2); PPBES, 3
samples from 2 stations (4).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 12-31 m, sand.
Yan, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 antenna 1 ordi-
nary, ventral setae widely spread or shifted apically.
Article 1 of antenna 2 not ensiform; article 3 with 2
setules; facial spines on article 4 in 2 or more rows;
article 5 short. Right mandibular incisor with 4+
NUMBER 245 389
teeth; molar not triturative, medium, pillow-
shaped, bearing 4 or more splayed semiarticulate
spines, usually not bearing fuzz; palpar hump me-
dium. Palp of maxilla 1 biarticulate; inner plate
with 3 setae. Setation of maxilla 2 ordinary. Inner
plate of maxillipeds ordinary; apex of palp article 3
weakly protuberant, dactyl elongate, apical nail dis-
tinct, medium. Gnathopods ordinary, small, similar,
article 5 of gnathopods 1-2 of ordinary length to
elongate, free, with or without eusirid attachment;
palms oblique, hands of gnathopods 1-2 ordinary
to elongate, poorly setose anteriorly. Article 5 of
pereopods 1-2 setose posteroproximally. Article 3 of
pereopod 3 of broad form, articles 4-5 of pereopods
3-4 broad to medium, article 2 of pereopods 3-4
not setose posteriorly; pereopod 5 ordinary, article 2
naked ventrally, article 3 ordinary, dactyl vestigial.
Epimera 1-2 lacking long posterior setae, without
midfacial setae above ventral facial ridge, epimeron
3 ordinary. Urosomite 1 generally naked; urosomite
3 without dorsal hook or special process. Peduncle
of uropod 1 without apicoventral spike, without
special enlarged apicolateral-medial spine, pedun-
ular apices of uropods 1-2 not combed, inner ramus
of uropod 1 with marginal spines in one row, some
rami continuously spinose to apex, or bearing acces-
sory nails, inner ramus of uropod 2 ordinary. Uro-
pod 3 ordinary to small, article 2 of outer ramus
carrying 2 medium apical setae. Telson ordinary,
with only one apical spine on each lobe plus set-
ules.
DESCRIPTION.?Rostrum fully developed, con-
stricted. [Fuzz on article 1 of antenna 1 in male un-
known, calceoli on male antennae 1-2 unknown.]
Prebuccal parts ordinary, poorly separated from
each other, upper lip dominant. Right lacinia mo-
bilis bifid or simple, thin; mandibular palp thin,
article 1 short to slightly elongate, article 2 without
outer setae, apex of article 3 oblique. Lower lip
bearing cones. Outer plate of maxilla 1 with 11
spines, one spine especially thickened. Inner plates
of maxilliped thick, ordinarily setose; outer plates
small. Coxae 2-4 without special anterodorsal
humps. All posterior spines on article 6 of pereo-
pods 1-2 thick and stiff, midapical spine or seta
present. Article 2 of pereopod 5 without facial
setae. Peduncle of uropod 1 with dorsolateral spines
confined apically, medial spines confined apically;
peduncle of uropod 2 with only one medial spine
or setule confined apically. Peduncle of uropod 3
lacking extra subapical setae or spines. Telson with
ordinary pair of midlateral or dorsal setules on
each side.
TYPE-SPECIES.?Yan tiendi, new species.
COMPOSITION.?Yan errichus, new species.
RELATIONSHIP.?The reduced dactyl of pereopod
5 appears to be a character of degree sufficient to
separate these two species from the pool of species
in Birubius. The two species of Yan also have
slightly shortened second articles of antenna 1 and
fifth articles of antenna 2 and have slightly reduced
rami of uropod 3. The type-species, Y. tiendi, is
closer to the birubiun root than Y. errichus which
bears heavily modified gnathopods. In other re-
spects, Y. tiendi has affinities with B. gallangus (10),
a species known only for the male. The male of Y.
tiendi is unknown. The two species have numerous
differences and are therefore not opposite sexes of
the same species. Yan tiendi differs from B. gallan-
gus in the presence of setae on coxa 4, the presence
of an accessory apical nail on the outer ramus of
uropod 2, the absence of a midfacial ridge on pereo-
pod 3, the presence of one (not zero) spine on the
inner plate of the maxilliped, the absence of a large
apicomedial spine on the peduncle of uropod 2, a
poorly setose midapex on article 4 of pereopods 1-2,
broader plates of maxilla 2, the absence of a third
main set of facial spines on article 4 of antenna
2, much sharper teeth on the left lacinia mobilis
and a broader distal ramus on the right lacinia
mobilis.
Yan tiendi differs from B. muldarpus (9) in the
vestigial dactyl of pereopod 5, the presence of setae
on coxa 4, the accessory nail on the outer ramus of
uropod 2, reduced setation on epimeron 3, a short
rostrum, the presence of only one apical spine on
the inner plate of the maxilliped, and slightly
crowded or short row of setae on epimeron 2.
The presence of setae on coxa 4 suggests a point
of origin for Y. tiendi in the B. lorus-B. nammul-
dus (2-3) species group but Y. tiendi differs from
those species in the reduction of setae on epimeron
3, the accessory nail on uropod 2, the presence of
only one main apical spine on the inner plate of
the maxilliped, the relatively short gnathopodal
wrists, and the shape of coxa 4, which in those
species is somewhat more trapezoidal than in Y.
tiendi. Birubius lorus is also characteristized by
numerous ventrolateral setae on urosomite 1 and
uncrowded, numerous setae on epimeron 2.
390 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Key to the Species of Yan
Gnathopods ordinary, hands about half as wide as long, outer ramus of uropod 2 with accessory
nail, right lacinia mobilis bifid Y. tirndi, new species
Gnathopods thin, hands about one third as wide as long, outer ramus of uropod 2 lacking acces-
sory nail, right lacinia mobilis simple Y. errichus, new species
Yan tiendi, new species
FIGURES 199, 200
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about 65
percent of length; rostrum constricted, broad, short,
exceeding apex of article 1 on antenna 1. Eyes me-
dium-large, largely occluded with pigment. Article
1 on peduncle of antenna 1 about 1.1 times as long
as wide, about 2.1 times as wide as article 2, ventral
margin with about 9 sctules, unproduced dorsal
apex with one setule; article 2 about 0.9 times as
long as article 1, with ventral cycle of 4 setae; pri-
mary flagellum with 7 articles, about 0.9 times as
long as peduncle, bearing long aesthetascs; acces-
sory flagellum with 5 articles; spine formula on
article 4 of antenna 2 = 1-3-4, dorsal margin with
notch bearing 2 setae, ventral margin with 3 groups
of 1-3 long to medium setae, one ventrodistal long
spine; article 5 about 0.7 times as long as article 4,
facial spine formula = 1-2, dorsal margin naked,
ventral margin with 2 sets of setae, 2 ventrodistal
medium spines; flagellum about 1.2 times as long
as articles 4-5 of peduncle combined, with 7 arti-
cles. Mandibles with strong palpar hump; right in-
cisor with 3 teeth, one notch, and one accessory
tooth; left incisor with 5 humps in 2 branches;
right lacinia mobilis bifid, distal branch little
shorter than proximal, flabellate, subbifid, proxi-
mal branch simple, weakly pointed, with marginal
denticles; left lacinia mobilis with 4 teeth; right
rakers 5, plus 2 rudimentaries; left rakers 5; molar
in form of bulbous hump, right and left molars
with 6 primarily long spines plus one short spine
strongly disjunct, plume absent; palp article 1
weakly elongate, article 2 with 2 short inner setae,
article 3 about 1.1 times as long as article 2,
oblique, apex with 5 spine-setae, basofacial form-
ule = 0. Inner plate of maxilla 1 thin, bearing one
long apical pluseta and 2 apicolateral much shorter
setae; palp article 2 with 3 apicalmedial marginal
spines and 3 submarginal setae. Plates of maxilla 2
extending subequally, of subequal width, outer
with 3 apicolateral setae, inner with 2 medial
setae. Inner plate of maxiliiped with one apical
spine, 2 apicofacial setae, 3 medial setae; outer
plate with 3 medial and apical spines, no apico-
lateral setae; palp articles 1-2 each with one apico-
lateral seta, medial margin of article 2 moderately
setose, article 3 weakly protuberant, with 2 facial
setae, one lateral seta, nail of article 1 medium,
with 2 accessory sctules. Coxa 1 scarcely expanded
(listally. anterior margin almost straight; main ven-
tral setae of coxae 1-4 - 5-5-5-2, posteriormost
seta of coxae 1-3 shortened; anterior and posterior
margins of coxa 4 divergent, posterior margin al-
most straight, posterodorsal corner sharply rounded,
posterodorsal margin long, concave, undulant,
width-length ratio of coxa 4 = 5:6. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 2-3-3-2, others not counted. Gnathopods
ordinary; width ratios of articles 5-6 on gnathopods
1-2 = 26:32 and 25:32, length ratios = 64:64 and
58:60: palmar humps ordinary, palms oblique;
article 5 of gnathopod 1 elongate, ovate, posterior
margin rounded, but almost flat; article 5 of gna-
thopod 2 weakly elongate, ovate, posterior margin
rounded. Pereopods 1-2 similar; facial setae form-
ula on article 4 = 1-1, on article 5 = 3-2; main
spine of article 5 extending to M. 75 on article 6,
article 5 lacking proximoposterior spines; spine
formula of article 6 = 3 + 3 plus middistal seta,
some spines especially short; acclivity on inner
margin of dactyls of pereopods 1-2 obsolescent,
emergent setule long, midfacial pluseta very long,
highly distad. Coxae 5-7 posteroventral setule form-
ula = 1-1-0. Articles 4?5 of pereopods 3-4 broad
to medium, facial spine rows sparse, facial ridge
formula on article 2 of pereopods 3?5 = 0-2-2;
width ratios of articles 2, 4, 5, 6 of pereopod 3 =
52:42:38:17, of pereopod 4 = 68:41:26:13, of pereo-
pod 5 = 82:18:15:8, length ratios of pereopod
3 = 74:25:35:40, of pereopod 4 = 88:45:40:47, of
pereopod 5 = 108:24:15:23; article 2 of pereopod
5 almost reaching middle of article 5, dactyl
vestigial; medial apex of article 6 finely combed
NUMBER ?45 391
FIGURE 199.?Yan tiendi, new species, holotype, female "a," 2.47 mm.
and bearing 3 digital processes. Posteroventral
corner of epimeron 1 rounded-subquadrate, pos-
terior margin convex, anteroventral margin with
2 medium setae, posteroventral face with one long
seta, facial ridge weak; posteroventral corner of
epimeron 2 rounded-subquadrate, posterior mar-
gin weakly convex, facial setae = 4; posteroventral
corner of epimeron 3 weakly protuberant, with set-
ule sinus, small tooth, posterior margin weakly un-
dulant, with one setule notch, face with oblique
horizontal row of 2 setal-spines near ventral-poste-
rior margin. Urosomite 1 naked, articulation line in-
complete; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1-2 with articulate enlarged
392 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
FIGURE 200.?Yan tiendi, new species, holotype, female "a," 2.47 mm.
NUMBER 245 393
apical nails, outer rami of uropods 1-2 with one
accessory nail, inner ramus of uropod 1 with 2 ac-
cessory nails, outer ramus of uropod 1 with 3 dorsal
spines, inner with one, outer ramus of uropod 2
with one dorsal spine, inner with one dorsomedial
spine; peduncle of uropod 1 with 2 apicolateral
spines and 2 basofacial setae, medially with 2 mar-
ginal spines, apicalmost enlarged; peduncle of uro-
pod 2 with 4 dorsal spines, medially with one apical
setule. Peduncle of uropod 3 with 2 ventral spines,
dorsally with one lateral spine, one medial spine;
rami feminine, inner extending to M. 80 on article
1 of outer ramus, apex with 2 long setae, medial
and lateral margins naked, article 2 of outer ramus
short, 0.22, bearing 2 long setae, medial margin of
article 1 naked, lateral margin with 2 acclivities,
spine formula = 1-2-2, setal formula = 1-0-0.
Telson ordinary, length-width ratio = 15:14, not
fully cleft, each apex of medium width, rounded-
truncate, lateral acclivity shallow, weak, bearing
ordinary lateral setule, spine next medial longer
than setule, midlateral setules diverse. Cuticle with
ordinary bulbar setules.
DESCRIPTION OF JUVENILE (juveniles, 1.36-2.00
mm).?Eyes small to very small. Spine formula on
article 4 of antenna 2 either 4-3-1 or 3-3-1; article
5 with 2 or 1 facial spines. Setae of coxae 1-3 =
3-3-3. Epimeron 1 with one anteroventral seta and
0-1 posterofacial seta; epimeron 2 with 4, 3, or 2
facial setae; epimeron 3 with only one posterofacial
seta and one posterior setule. Peduncle of uropod 1
with 1-2 basofacial setae, peduncle with 2 or 1
apicolateral spines, outer ramus with 2 or 1 dorsal
spines, apex with nail and one accessory spine, in-
ner ramus with 0-1 dorsal spine, apex with nail
and 2 accessory spines; peduncle of uropod 2 with
2-3 spines, outer ramus lacking dorsal spines, apex
with nail and one accessory spine, inner ramus with
one or no dorsal spine, apex with nail and 0?1
accessory spine, this accessory spine occurring in 1.36
mm juvenile and apparently representing future
dorsal spine, similar situation true for inner ramus
of uropod 1 on both 1.36 and 2.00 mm juveniles.
Uropod 3 with 0-1 lateral acclivity and 0-1 lateral
spine.
ILLUSTRATIONS.?Outer plate of maxilla 1 drawn
obliquely; maxillipedal palp reduced in magnifica-
tion in comparison to inner and outer plates; pereo-
pod 3 illustrated from medial view but lateral
ridges definitely absent; pereopodal dactyl illus-
trated from oblique view but acclivity obsolescent
when turned to precisely lateral view; outer ramus
of uropod 2 definitely with 2 apical setae, one miss-
ing in illustration.
HOLOTYPE.?WAM, female, "a," 2.47 mm, brood-
pouch with 3 large eggs.
TYPE LOCALITY.?JLB AUS 11, 30 Sep 1968,
Middleton Beach, Albany, Western Australia, inter-
tidal wash of algae and rocks.
VOUCHER MATERIAL.?Type-locality, juvenile
"b," 2.00 mm; Slack-Smith 1, juvenile "c," 1.36
mm.
RELATIONSHIP.?See generic remarks and rela-
tionships of Yan errichus.
MATERIAL.?JLB AUS and Slack-Smith, 3 sam-
pies (7).
DISTRIBUTION.?Western Australia, Albany, inter-
tidal.
Yan errichus, new species
FIGURES 201, 202
DESCRIPTION OF MALE (young).?Head about 18
percent of total body length, greatest width about
72 percent of length; rostrum weakly constricted,
broad, short, reaching apex of article 1 on antenna
1. Eyes medium, largely occluded with pigment.
Article 1 on peduncle of antenna 1 about 1.1 times
as long as wide, about twice as wide as article 2,
ventral margin with 5-6 setules, weakly produced
dorsal apex with 3 setules; article 2 about 0.9 times
as long as article 1, with apicoventral cycle of 3
setae; primary flagellum with 7 articles, about 0.85
times as long as peduncle, bearing several aesthe-
tascs; accessory flagellum with 4 articles. Spine
formula on article 4 of antenna 2 = 1-3-3-1, dorsal
margin with notch bearing 2 setae, ventral margin
with 2 groups of 2 long to short setae, one ventro-
distal medium spine; article 5 about 0.75 times as
long as article 4, facial spine formula = 1-2, dorsal
margin bearing 2 setules, ventral margin with one
setule and seta-setule pair, 2 ventrodistal short to
medium spines; flagellum about 1.6 times as long
as articles 4-5 of peduncle combined, with 9 arti-
cles. Mandibles with weak palpar hump; right in-
cisor with 3 teeth and accessory notch; left incisor
with 2 branches in 5 humps; right lacinia mobilis
simple, narrow; left lacinia mobilis with 4 teeth;
right rakers 3 plus one rudimentary; left rakers 4
394 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
M ' ^ / S
FIGURE 201.?Yan errichus, new species, holotype, male "a," 2.74 mm.
plus 2 rudimentaries; molar in form of elongate
plaque, each molar with 5 short to medium spines
plus one short spine strongly disjunct, no plumes;
palp article 1 slightly elongate, article 2 with 2
short inner setae, article 3 about 1.15 times as long
as article 2, oblique apex with 4 spine-setae, baso-
facial formula = 0. Inner plate of maxilla 1 ordi-
nary, bearing one short apicomedial seta and 2
apicolateral much shorter setae; palp article 2 with
3 apicalmedial marginal spines and 2 submarginal
setae. Plates of maxilla 2 extending subequally, of
equal width, outer with 2 apicolateral setae, inner
with no medial setae. Inner plate of maxilliped
with one large thick apical spine, 2 apicofacial
setae, 3 medial setae; outer plate with 4 medial and
apical spines, no apicolateral setae; palp article 1
lacking apicolateral seta, article 2 with one apico-
lateral seta, medial margin of article 2 moderately
setose, article 3 weakly protuberant, with 2 facial
setae, no lateral setae, nail of article 4 medium,
with 2 accessory setules. Coxa 1 scarcely expanded
distally, anterior margin weakly convex; main
NUMBER 245 395
W
FIGURE 202.?Yan errichus, new species, holotype, male "a," 2.74 mm.
3% SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ventral setae of coxae 1-4 = 4-4-5-3, posteriormost
seta of coxae 1-3 shortest; anterior and posterior
margins of coxa 4 strongly divergent, posterior
margin very oblique, posterodorsal corner rounded,
posterodorsal margin short, weakly concave, width-
length ratio of coxa 4 = 16:17. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 1-1-2-2, short anteriors = 1-1-1-1, long
anteriors = 2-3-0-0, no others. Gnathopods very
thin; width ratios of articles 5-6 on gnathopods
1-2 = 21:25 and 21:29, length ratios = 59:68 and
51:65; palmar humps ordinary, sharp, palms
oblique; article 5 of gnathopod 1 elongate, ovate,
posterior margin rounded; article 5 of gnathopod
2 elongate, ovate, posterior margin rounded. Pereo-
pods 1-2 similar, facial setae formula on article
4 = 2 and 2, on article 5 = 2 and 2; main spine of
article 5 extending to M. 90 on article 6, article 5
with no proximoposterior spines; spine formula of
article 6 = 3 + 2 plus middistal seta on both pairs,
spines short; acclivity on inner margin of dactyls of
pereopods 1-2 sharp, produced as tooth, emergent
setule long, midfacial pluseta ordinary, highly
distad. Coxae 5-7 posteroventral setule formula =
1-1-1. Articles 4-5 of pereopods 3-4 of ordinary
width, facial spine rows sparse, facial ridge formula
on article 2 of pereopods 3-5 = 0-2-2; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 45:38:34:
16, of pereopod 4 = 58:35:25:13, of pereopod
5 = 75:17:14:8, length ratios of pereopod 3 =
67:29:29:37, of pereopod 4 = 80:40:34:41, of pe-
reopod 5=100:22:20:21; article 2 of pereopod
5 reaching apex of article 4, dactyl apparently
absent; medial apex of article 6 bearing 3 digital
processes. Posteroventral corner of epimeron 1
rounded, posterior margin deeply convex, corner
with setule, anteroventral margin with one medium
seta, posteroventral face with one long seta, ridge
absent or faint; posteroventral corner of epimeron
2 rounded, posterior margin strongly convex, facial
setae = 5, ridge absent; posteroventral corner of
epimeron 3 with small rounded tooth, posterior
margin weakly convex, face with horizontal row of
2 setae; epimera 1-3 with setule on posterodorsal
margin set in weak notch. Urosomite 1 with articu-
lation line incomplete; urosomite 3 weakly pro-
tuberant dorsally. Rami of uropods 1-2 with articu-
late, enlarged apical nails, inner ramus of uropod
1 with two accessory nails, outer ramus of uropod 1
with 2 dorsal spines, inner with one, outer ramus
of uropod 2 with one dorsal spine, inner with none;
peduncle of uropod 1 with 2 apicolateial spines,
and one basofacial seta, medially with one apical
enlarged spine; peduncle of uropod 2 with 3 dorsal
spines, medially with one medium apical spine;
apicolateral corners on peduncles of urojxxls 1-2
lacking comb. Peduncle of uropod 3 with 5 ventral
spines, dorsally with one lateral spine, one medial
spine; rami submasculinc, inner extending to M.
120(!) on article 1 of outer ramus, apex with 2 setae,
medial margin with 2 setae, article 2 of outer ramus
short, 0.11, bearing 2 medium setae, apitomedial
margin of article 1 with one seta, medial margin
with 2 other setae, lateral margin with 3 acclivities,
spine formula 1-1-1-2, setal formula 0. Tclson
ordinary, length-width ratio = 6:5, not fully cleft,
each aj)t'x wide, rounded, lateral acclivity obsoles-
cent, bcai ing short lateral setule, spine next medial
of length equal to setule, midlatcral setulcs ordi-
nary. Cuticle with ordinary bulbar setules of varying
sizes mixed with pipes, surface bearing fine stria-
tions in form of linear fingerprint pattern, emergent
setules branched.
ILLUSTRATIONS.?Upper lip as shown for Y. tiendi,
but ventral margin straight; epistome-upper lip
from lateral view as in Y. tiendi but lacking antero-
ventral hump on anterior margin; lower lip with
one cone on each lobe, mandibular lobes short and
broad; general appearance of maxilla 2 as in
Birubius karobrani, but inner lobe with 3 facial
setae, 4 retrofacial setae and 2 other distal setae;
dactyl of pereopod 5 apparently absent, socket on
article 6 not present.
HOLOTYPE.?AM, male "a," 2.74 mm. Unique.
TYPE-LOCALITY.?AM P. 18214, 16 Nov 1970,
Long Reef, Collaroy, New South Wales, Australia.
RELATIONSHIP.?This species differs from Y.
tiendi in the much thinner gnathopods, the simple
right lacinia mobilis, and the absence of an acces-
sory apical spine on the outer ramus of uropod 2;
various distinctions in spination and setation can
also be seen in the illustrations.
MATERIAL.?AM, one sample (1).
DISTRIBUTION.?New South Wales, Long Reef,
Collaroy.
Tickalerus, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
NUMBER 24S 397
elongate, main setae midventral. Article 1 of an-
tenna 2 not ensiform; article 3 with 2 setules; facial
spines on article 4 in 3 rows; article 5 ordinary in
size. Right mandibular incisor with 3-4 teeth (in-
terpretive); molar not triturative, ordinary, med-
ium, pillow-shaped, bearing 4 or more splayed,
semiarticulate spines, usually not bearing fuzz;
palpar hump small. Palp of maxilla 1 biarticulate;
inner plate with 4 setae. Setation of maxilla 2 ordi-
nary. Inner plate of maxillipeds ordinary; apex of
palp article 3 weakly protuberant, dactyl elongate,
apical nail distinct, medium. Gnathopods ordinary,
small, similar; article 5 of gnathopods 1-2 free, of
ordinary length, or elongate on gnathopod 1, with-
out eusirid attachment; palms oblique, hands of
gnathopods 1-2 ordinary, ovatorectangular, poorly
setose anteriorly. Article 2 of percojxxi 3 of broad
form, articles 4-5 of jjereopods 3-4 broad, article 2
of pereopods 3-4 not setose posteriorly; pereopod 5
ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Epimera 1-2 lacking long pos-
terior setae, without midfacial setae above ventral
facial ridge; epimeron 3 ordinary. Urosomite 1 gen-
erally naked; urosomite 3 with dorsal hook or spe-
cial process. Peduncle of uropod 1 normally elong-
ate, without apicoventral spike, without special
enlarged apicolateral or medial spine; peduncular
apices of uropods 1-2 not combed; inner ramus of
uropod 1 with marginal spines in one row, no rami
continuously spinose to apex, inner ramus of uro-
pod 2 ordinary. Uropod 3 somewhat reduced, rami
scarcely longer than peduncle, article 2 of outer
ramus carrying 2 long apical setae. Telson ordinary,
with only one apical spine on each lobe.
DESCRIPTION.?Rostrum fully developed, con-
stricted, small. [Fuzz on article 1 of antenna 1 in
male and calceoli on male primary flagellum of
antenna 1 and antenna 2 unknown.] Prebuccal
parts ordinary, poorly separated from each other,
upper lip dominant. Right lacinia mobilis bifid,
flabellate; mandibular palp thin, article 1 slightly
elongate, article 2 without outer setae, apex of
article 3 oblique. Lower lip bearing cones. Outer
plate of maxilla 1 with 11 spines, one spine espe-
cially thickened. Inner plates of maxilliped thick,
ordinarily setose. Coxae 2-4 without special antero-
dorsal hump. All posterior spines on article 6 of
pereopods 1-2 thick and stiff, midapical seta vestig-
ial. Article 2 of pereopod 5 without facial setae.
Peduncle of uropod 1 with dorsolateral spines con-
fined apically; medial spines confined apically;
peduncle of uropod 2 with only one medial spine
or setule confined apically. Peduncle of uropod 3
lacking extra subapical setae or spines. Telson with
ordinary pair of midlateral or dorsal setules on each
side.
TYPE-SPECIES.?Tickalerus birubi, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This monotypic genus differs
from Birubius in the enlarged and sharp dorsal
process on urosomite 3, the shortness of the rami on
uropod 3, and in the development of 3 distinct sets
of dorsal setae on article 4 of antenna 2. These
characters are seen rudimentarily in several species
of Birubius. For example, B. apart (6) has a weakly
thrust urosomite 3, B. gambodeni (20) has super-
numerary dorsal setae on article 4 of antenna 2, and
several species show the rudiments of shortness in
rami on uropod 3.
Two other species similar to Tickalerus birubi
have been segregated into the following genus,
Kulgaphoxus, but, as explained therein, they differ
from Tickalerus by an unevenly mixed set of
generic characters. This implies they are polyphy-
letic. One of these characters, found only in one
species, is the posteroventral bevelment of article 2
on pereopod 5, an attribute unknown in Birubius.
The 3 species share similarities in the presence of
posterior straw-setae on article 2 of several members
of gnathopods 1-2 and pereopods 1-2, but unlike
Tickalerus, the two species of Kulgaphoxus have
accessory nails on the inner rami of both uropods
1-2. The two genera also share, at least in part, by
discounting K. cadgeeus, other characters suggest-
ing that they have a common ancestor: the presence
of more than 2 posteroproximal spines on article 5
of pereopods 1-2; the extreme reduction of the
middistal seta on article 6 of those pereopods; the
loss of plumosities on the midfacial dactylar seta of
those pereopods; and the reduction of uropod 3,
with elongation of the peduncle.
The single species so far known for Tickalerus
differs in many other characters from either species
of Kulgaphoxus but, in general, appears to be the
more primitive grade of evolution in terms of
greater diversity in spine rows on article 4 of an-
tenna 2, the more primitive molar (similar to
Birubius), and the shorter article 5 of gnathopod 2.
But Tickalerus also differs in characters about
which the level of primitiveness is unknown, such
398 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
as the shape of coxa 4. Illustrations and descrip-
tions of the following parts also should be com-
pared: prebuccal, mandibular palp, right lacinia
mobilis, palp of maxilla 1, dactyl of pereopod 5,
uropod 3, telson, coxa 1 expansion, coxa 3, and the
outer ramus of uropod 1.
Tickalerus bbrubi, new species
FIGURES 203-205
DESCRIPTION OF FEMALE.?Head about 17 percent
of total body length, greatest width about 75 per-
cent of length; rostrum constricted, narrow, short,
exceeding apex of article 1 on antenna 1. Eyes med-
ium, clear of pigment, ommatidia especially small.
Article 1 on peduncle of antenna 1 about 1.4 times
as long as wide, about 2.2 times as wide as article 2,
ventral margin with about 13 setules, weakly pro-
duced dorsal apex with one setule, article 2 about
0.85 times as long as article 1, with ventral cycle of
11 setae; primary flagellum with 13 articles, about
0.75 times as long as peduncle, bearing several long
aesthetascs; accessory flagellum with 10 articles.
Spine formula on article 4 of antenna 2 = 1-3-4-6,
dorsal margin with 3 notches bearing 3 setae each,
ventral margin with 7-8 groups of 2-4 long to short
setae, one ventrodistal long spine; article 5 about
0.7 times as long as article 4, facial spine formula =
1-2-2, dorsal margin naked, ventral margin with 7
sets of 1-3 long to short setae, 2 ventrodistal long to
medium spines; flagellum about 1.1 times as long as
articles 4-5 of peduncle combined, with 14 articles.
Mandibles with weak palpar hump; right incisor
with 3 teeth and notch; left incisor with 5 humps
in 2 branches; right lacinia mobilis bifid, distal
branch little shorter than proximal, flabellate,
broad, subbifid, denticulate, with weak facial
humps, proximal branch simple, pointed, with
marginal denticles and facial humps; left lacinia
mobilis with 4 teeth; right rakers 12; left rakers 11;
molar in form of bulbous hump, right molar with
10 primarily long spines plus one short spine
strongly disjunct, left molar with 7 primarily long
spines plus one short spine strongly disjunct; palp
article 1 slightly elongate, article 2 with one med-
ium inner apical seta and 2 other shorter inner
setae, article 1 with 2 short setae, article 3 about
1.1 times as long as article 2, oblique apex with 12
spine-setae, basofacial formula = 3-5. Inner plate
of maxilla 1 elongate, thin, bearing one long apical
pluseta, one shorter similar apicomedial seta, 2
apicolateral much shorter setae; palp article 2 with
4 apicalmedial marginal spines and 4 submarginal
setae. Plates of maxilla 2 extending equally, outer
broader than inner, with 6 apicolateral setae, inner
with 3 medial setae. Inner plate of maxilliped with
2 large, thick apical spines, 3 apicofacial setae, 5
medial setae; outer plate with 8 medial and apical
spines, 3 apicolateral setae; palp article 1 with api-
colateral seta, article 2 with 2 groups of 3 apicolat-
eral setae, medial margin of article 2 moderately
setose, article 3 with 6 facial setae, set of 3 lateral
setae, nail of article 4 medium-short, with 2 acces-
sory setules. Coxa 1 scarcely expanded distally,
anterior margin weakly concave, main ventral setae
of coxae 1-4 = 12-10-12-0. posterior most seta of
coxae 1-3 shortened; anterior and posterior margins
of coxa -\ parallel, posterior margin straight, pos-
terodorsal corner sharp, posterodorsal margin short,
concave, width-length ratio of coxa 4 = 2:3(1).
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = (12-13)-(16-19)-13-9, short
posteriors = 0-2-3-4, long anteriors = (6?7)?18?
1-0, short anteriors = (8-10)-8-11-(4-6). Gnatho-
pods ordinary; width ratios of articles 5-6 on
gnathopods 1-2 = 25:34 and 27:37, length ratios
= 66:57 and 59:58; palmar humps ordinary to
large, palms oblique; article 5 of gnathopod 1
elongate, ovate, posterior margin flat; article 5 of
gnathopod 2 ovate, posterior margin rounded, al-
most lobate. Pereopods 1-2 similar, facial setae
formula on article 4 = 8-9 and 5-6, on article 5 =
8 and 10; main spine of article 5 extending to M.
85 on article 6, article 5 with 4 and 3 proximopos-
terior spines; spine formula of article 6 = 7 + 9 or
6 + 9 or 6 + 8 and 7 + 1 0 plus vestigial middistal
seta, some spines especially long; acclivity on inner
margin of dactyls of pereopods 1-2 weak but pro-
duced as tooth, emergent setule short, midfadal
pluseta absent. Coxae 5-7 posteroventral seta for-
mula = 9-10-6. Articles 4-5 of pereopods 3-4
broad; facial spine rows dense, facial ridge formula
on article 2 of pereopods 3-5 = 0-2-2, anterior on
pereopod 5 short; width ratios of articles 2, 4, 5,6 of
pereopod 3 = 50:60:44:18, of pereopod 4 = 80:
57:42:14, of pereopod 5 = 92:24:18:8, length
ratios of pereopod 3 = 86:50:46:39, of pereopod
4 = 107:70:50:55, of pereopod 5 = 120:27:23:23;
article 2 of pereopod 5 exceeding apex of article
NUMBER 245 399
FIGURE 203.?Tickalerus birubi, new species, holotype female "a," 8.1 mm (c = juvenile female
"c," 6.30 mm; v ? male "v," 6.6 mm; K ? spine in molar).
400 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 204.?Tickalerus birubi, new species, holotype, female "a," 8.1 mm (v ? male "v,
6.6 mm).
NUMBER 245 401
FIGURE 205.?Tickalerus birubi, new species, holotype, female "a," 8.1 mm (v = male "v,1
6.6 mm).
4; medial apex of article 6 not combed, bearing
6 digital processes. Posteroventral corner of epi-
meron 1 rounded, protuberant, posterior margin
convex, corner with setule, anteroventral margin
with 8 long to medium setae, posteroventral face
with 6 long setae, posterior most pair set verti-
cally; posteroventral corner of epimeron 2 rounded,
weakly protuberant, with setule sinus, posterior
margin weakly convex, with setule notch, facial
setae =12, posteriormost pair set vertically, upper
member above facial ridge; posteroventral corner
of epimeron 3 rounded, weakly protuberant, with
setule sinus, posterior margin straight, with 3
setule notches, ventral margin naked, midface
with oblique row of 6 setae near posterior mar-
gin; epimeron 3 with setule on posterodorsal
margin set in weak notch. Urosomite 1 naked,
articulation line almost complete; urosomite 3
with large hook dorsally. Rami of uropods 1-2
with articulate enlarged apical nails, outer ramus of
402 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
uropod 1 with 7 dorsal spines, inner with one, outer
ramus of uropod 2 with 4 dorsal spines, inner with
one dorsomedial spine; peduncle of uropod 1 with
6 apicolateral spines and one basofacial seta, med-
ially with 2 marginal setae and spines and apical
enlarged spine; peduncle of uropod 2 with 8 dorsal
spines, medially with one small apical spine.
Peduncle of uropod 3 with 6 ventral spines, dorsally
with one lateral spine, one medial spine and 3
setules; rami masculine, inner extending to M.
100+ on article 1 of outer ramus, apex with 3 setae,
medial and lateral margins setose, article 2 of outer
ramus short, 0.12, bearing 2 long setae, apicomed-
ial margin of article 1 setose, lateral margin with 4
acclivities, spine formula = 2-1-1-1-1, setal for-
mula - 1-2-2-2-2. Telson long, length-width ratio
= 31:26, not fully cleft, each apex wide, rounded,
lateral acclivity deep, broad, bearing short lateral
setule, spine next medial much longer than setulc,
with 1-2 medial setules, midlateral setules diverse.
Cuticle with enlarged bulbar setules, emergent
setules plumose.
DESCRIPTION OF MALE (very young).?Scarcely dis-
tinct from female in secondary sexual attributes
such as head, antenna 1, maxilla 1, coxae, pereo-
pods 3-4, uropods 1-3, epimera 1-2, uropod 3 with
fewer setae than in female because of youthfulness.
Eyes scarcely enlarged. Article 1 of antenna 1 lack-
ing medial fuzz. Spine formula on article 4 of an-
tenna 2 = 1-3-4-5, on article 5 = 1-2; flagellum
proliferate. Right lacinia mobilis somewhat more
complex than in female; right mandibles with 9
rakers, left with 10 + one rudimentary; basofacial
setae on article 3 of mandibular palp = 2 + 3.
Maxilla 2 with one fewer seta medially and apico-
laterally (2 and 5). Maxilliped less setose. Postero-
proximal spines on article 5 of pereopods 1-2 better
developed than in female (see illustration), dactyl
bearing midfacial nonplumose seta. Article 2 of
pereopod 5 slightly broader than in female (juve-
noid condition). Epimeron 3 with one anterofacial
seta, oblique row with 5 setae. Telson slightly
elongate.
OBSERVATIONS.?Prebuccal complex unproduced
anteriorly; pereopod 5 with tiny setules on anterior
margin of article 2; short posterior setae on article
2 of gnathopods 1-2 and pereopods 1-2 of variety
known as "straw."
ILLUSTRATIONS.?Known males scarcely distinct
from females, therefore head of male not illus-
trated; disjunct spine on molars difficult to inter-
pret owing to small size, transparency and tendency
to roll into arc fitting curve of molar; bulbar setules
so conspicuous as to be included on several illustra-
tions; pleonal view of holotype with apices of setae
on uropod 3 broken off, one spine on peduncle of
uropod 2 reconstructed; palp of maxilla 1 opposite
to that illustrated bearing one fewer apical seta;
short posterior seta on coxa 2 missing.
HOLOTYPE.?NMV, female "a." 8.1 mm.
TYPE LOCALITY??PBS 24N/5, 1 Mar 1965,
Western Port, Victoria, Australia, 9.1 m, muddy
sand.
VOUCHER MATERIAL.?CPBS 26N/4, male "v," 6.6
mm (illus.); CPBS 26N/2. juvenile female "c." 6.50
mm (illus.).
RELATIONSHIP.?See remarks after generic descrip-
tion.
MATERIAL.?CPBS, fi samples from 2 stations (8).
DISTRIBUTION.?Victoria, Western Port, 9.1-10.9
m, sand or muddy sand.
Kulgaphoxus, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially elongate, ventral setae narrowly spread
and proximal. Article 1 of antenna 2 not ensiform;
article 3 with 2 setules; facial spines on article 4 in
2 rows; article 5 short. Right mandibular incisor
with 3-4 teeth (interpretative); molar not tritura-
tive, ordinary, medium, truncate-conical, bearing 4
or more splayed, semiarticulate spines, not bearing
fuzz; palpar hump small. Palp of maxilla 1 biarticu-
late; inner plate with 4 setae. Setation of maxilla 2
ordinary. Inner plate of maxillipeds ordinary; apex
of palp article 3 weakly protuberant, dactyl elon-
gate, apical nail distinct, medium to elongate. Gna-
thopods ordinary, small, similar; article 5 of gna-
thopods 1-2 free, elongate, without eusirid attach-
ment; palms oblique, hands of gnathopods 1?2
ordinary, ovatorectangular, elongate, or broadened,
poorly setose anteriorly. Article 5 of pereopods 1-2
setose posteroproximally. Article 2 of pereopod 3 of
broad form, articles 4-5 of pereopods 3-4 broad,
article 2 of pereopods 3-4 not setose posteriorly;
pereopod 5 ordinary, article 2 naked or bearing one
slightly elongate seta ventrally, article 3 ordinary,
dactyl normal or vestigial. Urosomite 1 generally
naked except for sparse apicoventral setae or spines
NUMBER 245 403
near base of uropod 1; urosomite 3 with dorsal
hook, or special process. Peduncle of uropod 1
slightly shortened, without apicoventral spike, with-
out special enlarged apicomedial spine; peduncular
apices of uropods 1-2 not combed; inner ramus of
uropod 1 with marginal spines in one row, some
rami with accessory apical nails; inner ramus of
uropod 2 slightly shortened. Uropod 3 small, rami
shortened, article 2 of outer ramus carrying 2 long
apical setae. Telson ordinary to short, with 1-2
apical spines on each lobe plus setules. Epimera
1-2 lacking numerous long posterior setae, without
midfacial setae above ventral facial ridge, epimeron
3 ordinary.
DESCRIPTION.?Rostrum poorly developed, con-
stricted, small. Fuzz on article 1 of antenna 1 in
male present: calceoli on male primary flagellum of
antenna 1 present. Calceoli on article 5 of male
antenna 2 present; flagellum in male with calceoli.
Prebuccal parts ordinary, especially massive,
strongly distinct, either upper lip dominant or both
parts equally prominent. Right lacinia mobilis
bifid, flabellate; mandibular palp thin, article 1
slightly elongate, article 2 without outer setae, apex
of article 3 oblique. Lower lip bearing cones. Outer
plate of maxilla 1 with 11 spines, one spine espe-
cially thickened. Inner plates of maxilliped thick,
ordinarily setose. Coxae 2-4 without special antero-
dorsal humps. All posterior spines on article 6 of
pereopods 1-2 thick and stiff, midapical spine or
seta vestigial. Article 2 of pereopod 5 without facial
setae. Peduncle of uropod 1 with dorsolateral spines
confined apically, medial spines confined apically;
peduncle of uropod 2 with only one medial setule
confined apically. Peduncle of uropod 3 lacking
extra subapical setae or spines. Telson with ordi-
nary pair of midlateral or dorsal setules on each
side.
TYPE-SPECIES.?Kulgaphoxus borralus, new
species.
COMPOSITION.?Kulgaphoxus cadgeeus, new
species.
RELATIONSHIP.?Kulgaphoxus differs from Biru-
bius in the shortened rami of uropod 3 equal to the
peduncle in length, and in the enlarged dorsal
hook of urosomite 3.
Birubius apari (6) has the rudiments of a dorsal
process on urosomite 3 as in Kulgaphoxus and
shares many other similarities, such as in gnatho-
pods, narrow spread of setae on article 2 of antenna
1, the somewhat shortened rami of uropod 3 but
the inner ramus of uropod 2 in B. apari lacks ac-
cessory nails.
Kulgaphoxus differs from Tickalerus in numer-
ous characters among which are many not attrib-
uted generic status in Birubius. Kulgaphoxus and
Tickalerus, are, however, so divergent, that one
must assume for the time being that they had dif-
ferent origins out of Birubius. Indeed, the two
species now placed in Kulgaphoxus themselves may
not be congeneric.
Kulgaphoxus differs from Tickalerus in the
smaller rostrum, more strongly dominant epistome
(but not equally so in the 2 species of Kulga-
phoxus), the proximal placement of ventral setae
on article 2 of antenna 1, the presence of only one
dorsal set of setae on article 4 of antenna 2 (like
Birubius), the presence of accessory nails on the
inner rami of uropods 1-2, and the perfectly rec-
tangular coxa 4.
Somewhat more minor and variable characters
include the following: reduction in number of ven-
tral setae on article 2 of antenna 1, presence of only
one mid set of facial spines on article 5 of antenna
2, reduction in basofacial setae on article 3 of
mandibular palp, reduction in size of molar and
loss of disjunctness of one spine, less grossly devel-
oped right lacinia mobilis, reduction in facial and
lateral seate on article 3 of maxillipedal palp, more
elongate article 5 of gnathopod 2, presence of dorso-
posterior tooth on article 2 of pereopod 3, even
shorter rami of uropod 3, more arcuate coxa 3, re-
duced spination on outer ramus of uropod 1, and
enlargement of spines on uropod 3.
The two species of Kulgaphoxus mix together
characters partially shared with Ticklarus so that
the evolutionary flow is highly uneven; for exam-
ple, one or the other but not both species of Kul-
gaphoxus shares with Tickalerus the following
characters: the unusual bevel on article 2 of pereo-
pod 5, similarities in palp of maxilla 1, elongation
of article 6 on pereopods 3-4, well developed dactyl
of pereopod 5, unexpanded coxa 1, normally elon-
gate telson, and the presence of anterior setules on
article 2 of pereopod 5.
One or the other species of Kulgaphoxus also
bears many similarities to the genus Yan, among
them the essential presence of only 2 rows of facial
spines on article 4 of antenna 2, the somewhat re-
duced rostrum, shortened rami of uropod 3, degree
404 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
of vestigiality or absence of midapical seta on arti-
cle 6 of pereopods 1-2 and the presence of accessory
nails on the rami of uropods 1-2. However, as in
Kulgaphoxus, the species of Yan are diverse; for
example, one species of Yan has 2 full rows of facial
spines on article 4 of antenna 2 plus one spine in a
third row, and there are strong distinctions in place-
ment of accessory nails on uropods 1-2. Yan can be
separated distinctly from Kulgaphoxus only on the
absence of a dorsal process on urosomite 3. Both
species of Yan have a vestigial dactyl on pereopod
5 whereas this situation occurs only in one species
of Kulgaphoxus. Yan has a shorter article 2 of an-
tenna 1 with ventral setae either in the middle or
set distally. Both species of Yan have a much better
developed molar than does Kulgaphoxus, somewhat
similar to Tickalerus. Both species of Yan have long
ventral setae (2 only) on coxa 4 unlike Tickalerus
and Kulgaphoxus, but the shape of coxa 4 is alike
between Kulgaphoxus and Yan whereas it is per-
fectly rectangular in Tickalerus.
Kulgaphoxus borralus, new species
FIGURES 206-208
DESCRIPTION OF FEMALE.?Head about 15 per-
cent of total body length, greatest width about 95
percent of length; rostrum constricted, broad, short,
exceeding middle of article 1 on antenna 1. Eyes
medium, largely occluded with pigment. Article 1
on peduncle of antenna 1 about 1.4 times as long
as wide, about 2.7 times as wide as article 2, ventral
margin with 6-8 setules, weakly produced dorsal
apex with 2 setules; article 2 about 1.1 times as
long as article 1, with proximoventral cycle of 7-8
setae; primary flagellum with 8 articles, about 0.45
times as long-as peduncle, bearing several aesthe-
tascs; accessory flagellum with 8 articles. Spine
formula on article 4 of antenna 2 = 1-3-8 or 1-3-7,
dorsal margin with notch bearing 2 setae, ventral
margin with 7 groups of 2-3 long to short setae,
one ventrodistal long spine; article 5 about 0.7
times as long as article 4, facial spine formula =
1-1, dorsal margin naked, ventral margin with 4
sets of one long seta, 2 ventrodistal long spines, one
set facially; flagellum about 1.2 times as long as
articles 4-5 of peduncle combined, with 12 articles.
Mandibles with weak palpar hump; right incisor
with 3 teeth and notch; left incisor with 6 humps in
2 branches; right lacinia mobilis bifid, distal branch
little shorter than proximal, flabellate, denticulate,
proximal branch simple, pointed, with marginal
denticles and facial hump; left lacinia mobilis with
4 teeth plus one accessory tooth; right rakers 8; left
rakers 10; molar in form of short protrusion
demarcated mainly by spines, right molar with 8
primarily long spines, none disjunct, left molar
with 10 (3 hidden) primarily long spines, none div
junct; palp article 1 slightly elongate, article 2 with
one short inner apical seta and 3 other shorter
inner setae, article 3 about 1.05 times as long as
article 2, oblique a|>ex with 7 spine-setae, basofacial
formula = 0-1. Inner plate of maxilla 1 elongate,
thin, bearing one long apical pluscta, one longer
apicomedial seta. 2 upicolateral much shorter setae,
palp article 2 with 4 apicalmedial marginal spines
and 2 submarginal setae. Plates of maxilla 2 ex-
tending subequally, outer broader than inner, outer
with 3 apicolatcral setae, inner with 2 medial setae.
Inner plate of maxilliped with 2 large, thick apical
spines, 3 apicofacial thick setae, I medial thick
setae; outer plate with 5 medial and apical spines,
no apicolateral setae; palp article 1 with apicolat-
eral seta, article 2 with 2 apicolateral setae, medial
margin of article 2 weakly setose, article 3 with 2
facial setae, no lateral setae, nail of article 4 long,
with 2 accessory setules. Coxa 1 not expanded dis-
tally, anterior margin weakly convex; main ventral
setae of coxae \-$ = 7-8-8-0, posteriormost seta of
coxae 1-3 shortened; anterior and posterior margins
of coxa 4 divergent, posterior margin oblique,
straight, posterodorsal corner sharp, posterodorsal
margin short, almost straight, width-length ratio
of coxa 4 = 16:17. Long posterior setae on article
2 of gnathopods 1-2 and pereopods 1-2 = 5-7-7-4,
short posteriors = 0-0-1-1, long anteriors = 7-8-
0-0, short anteriors = 4-3-4-2. Gnathopods elong-
ate; width ratios of articles 5-6 on gnathopods 1-2
= 25:30 and 27:32, length ratios = 93:63 and
88:62, palmar humps ordinary, palms weakly
oblique; article 5 of gnathopods 1-2 elongate,
ovate, posterior margin flat, long. Pereopods 1-2
similar, facial setae formula on article 4 = 5-6 and
4, on article 5 = 5 and 4, main spine of article 5
extending to M. 80 on article 6, article 5 with 3
proximoposterior spines, spine formula of article
6 = 5 + 7 and 6 + 7 plus tiny vestigial middistal
seta, some spines especially long; acclivity on inner
margin of dactyls of pereopods 1-2 sharp, produced
NUMBER 245 405
s
FIGURE 206.?Kulgaphoxus borralus, new species, holotype, female "a," 4.90 mm.
406
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Gl
FIGURE 207.?Kulgaphoxus borralus, new species, holotype, female "a," 4.90 mm.
NUMBER 245 407
FIGURE 208.?Kulgaphoxus borralus, new species, holotype, female "a," 4.90 mm.
as tooth, setule fully immersed, midfacial seta non-
plumose (plumose on pereopods 3-5). Coxae 5?7
posteroventral seta formula = 6-13-16. Articles 4-5
of pereopods 3-4 broad; facial spine rows dense,
facial ridge formula on article 2 of pereopods 3-5
= 1-2-2; width ratios of articles 2, 4,5,6 of pereo-
pod 3 = 53:54:38:18, of pereopod 4 = 80:80:40:20,
of pereopod 5 = 87:20:15:7, length ratios of
pereopod 3 = 78:44:35:27, of pereopod 4 = 104:
60:40:40, of pereopod 5 = 110:22:22:22, article 2
of pereopod 5 almost reaching apex of article
5, article 2 of pereopod 3 with sharp tooth at
posterodorsal corner; medial apex on article 6 of
pereopod 5 bearing 4 weak digital processes, dactyl
vestigial. Posteroventral corner of epimeron 1
rounded, posterior margin deeply convex, antero-
ventral margin with 8 long to medium setae, pos-
teroventral face with 3 long setae, anterior pair set
vertically; posteroventral corner of epimeron 2
rounded, posterior margin strongly convex, facial
setae = 8, posteriormost pair set vertically; postero-
ventral corner of epimeron 3 weakly protuberant,
with 2 setule sinuses, posterior margin straight, with
2 setule notches, ventral marign naked, face with
oblique row of 10 setae in middle, epimeron 3 with
setule on posterodorsal margin set in deep notch.
Urosomite 1 naked, articulation line almost com-
plete; urosomite 3 with large hook dorsally. Rami
of uropods 1-2 with articulate enlarged apical nails,
inner rami of uropods 1-2 each with two accessory
408 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
nails, outer ramus of uropod 1 with 2 dorsal spines,
inner with one, outer ramus of uropod 2 with 2
dorsal spines, inner with none; peduncle of uropod
1 with 3 apicolateral spines and one basofacial
spinule, medially with 2 marginal setules and
apical enlarged spine; peduncle of uropod 2 with
7 dorsal spines, medially with one apical setule.
Peduncle of uropod 3 with 5 ventral spines, dorsally
with one lateral spine, one medial spine and setule;
rami submasculine, inner extending to M. 90 on
article 1 of outer ramus, apex with 2 setae, medial
and lateral margins setose, article 2 of outer ramus
short, 0.18, bearing 2 long setae, apicomedial
margin of article 1 with 2 setae, lateral margin with
3 acclivities, spine formula = 2-2-2-2, setal formula
= 0. Telson especially long, length-width ratio =
14:11, not fully cleft, each apex of medium width,
undulant, lateral acclivity narrow, bearing ordinary
lateral setule, spine next medial longer than setule,
second spine next medial as long as setule, mid-
lateral setules diverse. Cuticle with ordinary bulbar
setules mixed with occasional pipes and spicules,
emergent setules ordinary but weakly branched and
thorned.
OBSERVATIONS.?Prebuccal mass rounded anter-
iorly; female "w" with 3 dorsal spines on outer
ramus of uropod 1, 9 facial spines on epimeron 3
and 7 facial setae on epimeron 2; juvenile " j " spine
formula on article 4 of antenna 2 = 4-5, peduncle
of uropod 1 with 2 apicolateral spines, of uropod
2 with 5 dorsal spines, outer ramus of uropod 2
with only one dorsal spine, formula on article 1 on
outer ramus of uropod 3 = 2-2; epimeron 3 with 7
facial spines.
ILLUSTRATIONS.?View of prebuccal mass from
anterior aspect with upper lip not fully flattened;
lower lip with narrower illustrated outer lobe not
fully flattened, broader lobe fully flattened; apical
setae on primary flagellum of antenna 1 broken off;
illustration of pereopod 3 reduced to 83 percent of
size in pereopods 4-5; outer ramus of uropod 3 with
one spine missing on article 1. One side of telson
with spines missing.
HOLOTYPE.?NMV, female "a," 4.90 mm.
TYPE-LOCALITY.?PPBES 974/3, 13 Oct 1971,
Port Phillip Bay, Victoria, Australia, 5 m, sand.
VOUCHER MATERIAL.?PPBES 974/4, female "w,"
3.87 mm; WPBES 1725/3, juvenile " j , " 3.32 mm.
Male unknown.
RELATIONSHIP.?Kulgaphoxus borralus shares
with Tickalerus birubi the following characters not
shared with Kulgaphoxus cadgeeus; normal telson,
unexpanded coxa 1, unstunted article 6 of pereo-
pods 1-2, and the special bevel on article 2 of pereo-
pod 5.
MATERIAL.?PPBES, 2 samples from one station
(1): WPBES, 2 samples from one station (2).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 5-13 m, sand.
Kulgaphoxus cadgeeus, new species
FIGURES 209-212
DESCRIPTION OF FEMALE.?Head about 13 percent
of total body length, greatest width about 95 per-
cent of length; rostrum constricted, narrow, very
short, scarcely reaching along article 1 on antenna
1. Eyes medium, clear of pigment, ommatidia espe-
cially large. Article 1 on peduncle of antenna 1
about 1.5 times as long as wide, about twice as
wide as article 2, ventral margin with about 8
setules, weakly produced dorsal apex with 2 setules;
article 2 about 0.85 times as long as article 1, with
proximoventral cycle of 5 setae; primary flagellum
with 10 articles, about 0.8 times as long as peduncle,
bearing several long aesthetascs; accessory flagellum
with 9 articles. Spine formula on article 4 of an-
tenna 2 = 1-3-7, dorsal margin with notch bearing
2 setae, ventral margin with 3 groups of 2 long to
medium setae, one ventrodistal medium stout spine;
article 5 about 0.67 times as long as article 4, facial
spine formula = 1-3, dorsal margin naked, ventral
margin with 2 sets of 2 long to short setae, 2 ventro-
distal long to medium spines, one set facially; flagel-
lum about 1.45 times as long as article 4-5 of
peduncle combined, with 10-12 articles. Mandibles
with medium palpar hump; right incisor with 3
teeth and notch; left incisor with 5 humps in 2
branches; right lacinia mobilis bifid, distal branch
little shorter than proximal, distal branch flabellate,
denticulate, proximal branch simple, pointed, with
marginal denticles and facial humps; left lacinia
mobilis with 4 teeth; right rakers 8; left rakers 9;
molar in form of short protrusion or bulbous
humps demarcated mainly by spines, right molar
with 5 primarily long spines plus one short spine
weakly disjunct, left molar with 4 long spines, none
disjunct, each molar with plume; palp article 1
elongate, article 2 with one medium inner apical
NUMBER 245 409
s
FIGURE 209.?Kulgaphoxus cadgeeus, new species, holotype, female "a," 4.11 mm (b ? male "b,1
3.25 mm; w ? female "w," 5.3 mm).
410 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 210.?Kulgaphoxus cadgeeus, new species, holotype, female "a," 4.11 mm (b ? male
"b," 355 mm).
NUMBER 245
411
bGl
FIGURE 211.?Kulgaphoxus cadger us, new species, holotype, female "a," 4.11 mm (6 ? male
"b," 3.25 mm).
seta and one other shorter inner seta, article 3 about
0.8 times as long as article 2, oblique apex with 7
spine-setae, basofacial formula = 2-2 and 2-1. Inner
plate of maxilla 1 elongate, thin, bearing one long
apical pluseta, one shorter similar apicomedial seta,
2 apicolateral much shorter setae; palp article 2
with 6 apicalmedial spines and setae. Plates of
maxilla 2 extending equally, of equal breadth,
outer with 3 apicolateral setae, inner with 2 medial
setae. Inner plate of maxilliped with 2 large, thick
apical spines, 2 apicofacial setae, 4 medial setae;
outer plate with 6-7 medial and apical spines, one
apicolateral seta; palp article 1 with apicolateral
seta, article 2 with apicolateral seta, medial margin
of article 2 weakly setose, article 3 with 2 facial
setae, one lateral seta, nail of article 4 long, with 2
accessory setules. Coxa 1 strongly expanded distally,
anterior margin weakly concave; main ventral setae
of coxae 1-4 = 5-5-5-0, posteriormost seta of coxae
1-3 shortened; anterior and posterior margins of
coxa 4 strongly divergent, posterior margin very
oblique, almost straight, posterodorsal corner
412 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 212.?Kulgaphoxus cadgeeus, new species, holotype, female "a," 4.11 mm (b ? male "b,1
3.25 mm; w = female "w," 5.3 mm).
NUMBER 245 413
angular, posterodorsal margin short, convex,
width-length ratio of coxa 4 = 8:9. Long pos-
terior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 2- (3-4)-3-4, short posteriors =
(l-2)-(3-5)-5-7, long anteriors = 2-1-0-0, short
anteriors = 5-4-11-(8-11). Gnathopods ordinary;
width ratios of articles 5-6 on gnathopods 1-2 =
25:38 and 29:41, length ratios = 80:70 and 62:62;
palmar humps large, palms oblique; article 5
of gnathopod 1 elongate, ovate, posterior margin
rounded; article 5 of gnathopod 2 ovate, posterior
margin rounded, almost lobate. Pereopods 1-2 sim-
ilar; facial setae formula on article 4 = 3?4 and 4,
on article 5 = 3 and 5; main spine of article 5 ex-
tending to M. 83 on article 6, article 5 with 3-4 and
3-4 stout proximoposterior spines; spine formula of
article 6 ~ 3 + 4 and 4 + 4 plus vestigial middistal
seta, one spine especially long; acclivity on inner
margin of dactyls of pereopods 1-2 sharp, produced
as tooth, emergent setule short, midfacial pluseta
ordinary. Coxae 5-7 posteroventral setule formula
= 6-1-1. Articles 4-5 of pereopods 3-4 broad; facial
spine rows dense, facial ridge formula on article 2
of pereopods 3-5 = 1-2-2; width ratios of articles
2, 4, 5, 6, of pereopod 3 = 45:47:42:20, of pereo-
pod 4 = 62:49:39:16, of pereopod 5=82:17:
14:7, length ratios of pereopod 3 = 81:42:37:39,
of pereopod 4 = 88:54:40:47, of pereopod 5 =
110:22:22:23, article 2 of pereopod 5 almost reach-
ing apex of article 5, article 2 of pereopod 3
with large posterodorsal tooth; medial apex on arti-
cle 6 of pereopod 5 bearing 2 weak digital processes,
dactyl stunted. Postero ventral corner of epimeron 1
rounded, protuberant, with setule sinus, posterior
margin convex, anteroventral margin with 1-2
medium setae, postero ventral face with 2-3 long
setae, posterior pair set vertically; postero ventral
corner of epimeron 2 rounded, weakly protuberant,
with setule sinus, posterior margin convex, antero-
ventral margin with setules, facial setae = 5-8,
posteriormost or next posterior pair set vertically,
postero ventral corner of epimeron 3 weakly pro-
tuberant, with setule sinus, posterior margin
straight, with 1-2 setule notches, ventral margin
naked, face with oblique row of 5-11 setae near
posterior margin in middle; epimera 1-2 with setule
on posterodorsal margin set in weak notch. Urosom-
ite 1 with 1-2 midventral setules, articulation line
incomplete; urosomite 3 with hook dorsally. Rami
of uropods 1-2 with articulate enlarged apical nails,
inner rami of uropods 1-2 with two accessory nails,
outer ramus of uropod 1 with 2-3 dorsal spines,
inner with one, outer ramus of uropod 2 with 2
dorsal spines, inner with none; peduncle of uropod
1 with 2-3 apicolateral spines and 2 basofacial
setules, medially with 2 apical spines, one enlarged;
peduncle of uropod 2 with 5 dorsal spines, medially
naked. Peduncle of uropod 3 with 3 ventral spines,
dorsally with one lateral spine, one medial spine and
setule; rami feminine, short, inner extending to M.
75 on article 1 of outer ramus, apex with one seta,
medial and lateral margins naked, article 2 of outer
ramus ordinary, 0.25, bearing 2 medium to long
setae, apicomedial margin of article 1 naked, lateral
margin with 2 acclivities, spine formula = 2-2-2,
setal formula = 0. Telson short, length-width ratio
= 14:17, not fully cleft, each apex wide, subtrun-
cate, lateral acclivity deep, narrow, bearing ordi-
nary lateral setule, spine next medial much longer
than setule, midlateral setules diverse. Cuticle with
ordinary bulbar setules.
OBSERVATIONS (female).?Anterior setule of coxae
1-4 rather posteriad; article 4 of pereopod 3 with
posterior marginal scales (absent in male); article 2
of pereopod 5 with anterior setules; coxae 2-3
slightly tapering apically; occasional specimens
with setule proximal to spines on peduncle of uro-
pod 1 apicolaterally.
Female "w": A superfemale with following at-
tributes similar to holotype female: head, antenna
1, antenna 2, pereopods 3-4, left pereopod 5,
maxilla 1, maxilla 2, right mandibular palp [left
mandible missing], maxilliped, right mandible with
7 raker spines plus one rudimentary spine. This
female like holotype except for the following: right
lacinia mobilis (illustrated), right pereopod 5 article
2 slightly broader than in holotype, epimera 2-3
(illustrated), epimeron 1 with 4 long postero ventral
facial setae, outer ramus of uropod 1 with 4 spines,
peduncle of uropod 2 with 6 spines.
DESCRIPTION OF MALE.?Eyes enlarged. Article 2
of antenna 1 with 2 proximoventral setae; primary
flagellum with 9 articles, one calceolus each on arti-
cles 1-5; accessory flagellum with 9-10 articles.
Facial spine formula on article 4 of antenna 2 =
1-3-5 or 1-3-6, on article 5 = 2, latter with 3 sets
of dorsal male setae and 2 calceoli; flagellar formula
= 28, 2, 3, 5, 7 . . . 27. Right mandibular rakers 8,
left 9; right molar with 5 spines and one disjunct,
left with 5 alone; basofacial setal formula on article
414 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
3 of palp = 2 + 1 . Palps of maxilla 1 with 6 and 7
spines and setae. Coxa 4 broadened, ventral setal
formula of coxae 1^4 = 5-5-4-0. Gnathopod 1
thinner than in female. Article 5 of pereopods 1-2
with only 3 posteroproximal spines; setae of article
4 shortened; main spine rows on article 6 = 4 + 4.
Article 2 of only pereopod 5 narrower than in
female, ventral margin lacking elongate setule, arti-
cles 5-6 of pereopod 4 slightly thinner than in
female (see illustration). Epimera 1-3 broadened,
posterior margin of epimeron 3 not shortened; setal
formulas, epimeron 1 anteroventral = 3, postero-
ventral = 3, 2 of these set vertically, epimeron 2
facial = 5, epimeron 3 posterior = 3 setules, facial
= 11 (one aberrant sheathed), ventral = 0. Urosome
slender, hook enlarged. Spine formulas of uropods,
uropod 1 peduncle apicolateral = 2, basofacial = 2
setules, uropod 2 peduncle dorsal = 8, dorsal spines
on outer ramus of uropod 1 = 2, of uropod 2 = 2,
inner ramus of uropod 1 = 1, of uropod 2 = 0.
Ventral spines on peduncle of uropod 3 = 4, spine
formula on article 1 of outer ramus = 1-1-1-2-2,
setal formula = 0-1-1-1-1. Telson elongate, distal
spines shortened, each lobe with dorsal row of
denticles.
ILLUSTRATIONS.?Male epimeron 3 with one ab-
berrant seta bearing rooted sheath (aberrant);
female "w," illustration of epimeron 2 based on
reconstruction from right and left sides together,
both sides slightly damaged; several setae on
illustration of male uropod 3 truncated apically for
clarity.
HOLOTYPE.?NMV, female "a," 4.11 mm.
TYPE-LOCALITY.?CPBS 35S/5, 18 Mar 1965,
Western Port, Victoria, Australia, 15.1 m, sand.
VOUCHER MATERIAL.?CPBS 25N/2, female "w,"
5.13 mm (illus.); RHM, off Crib Point, 22 Oct 1971,
male "b," 3.25 mm (illus.); CPBS 34N/4, female
'%" 4.08 mm.
RELATIONSHIP.?Kulgaphoxus cadgeeus shares
with Tickalerus birubi the following characters not
shared with K. borralus: the presence of anterior
setules on article 2 of pereopod 5, the fully medial
spines on the palp of maxilla 1, the elongate article
6 of pereopods 3-4 and the well developed dactyl
of pereopod 5. Kulgaphoxus cadgeeus differs from
K. borralus in these additional characters: shorter
telson bearing only one main spine on each lobe,
the expanded coxa 1 and the smaller rostrum.
Unlike either K. borralus or T. birubi, Kulga-
phoxus cadgeeus lacks the special bevel on article
2 of pereopod 5, and has a set of 3 facial spines on
article 5 on antenna 2.
MATERIAL.?CPBS, 7 samples from 4 stations (9);
PPBES, 2 samples from 2 stations (6); RHM, one
sample (1).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 1.0-15.1 m and neritic, sand, coarse
sand, clayey sand.
Microphoxus J. L. Barnard
Microphoxus]. L. Barnard, 1960:291.
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female, article 2 of antenna 1
ordinary, ventral setae widely spread. [?Article I
of antenna 2 not ensiform; article 3 with 2 setules;
facial spines on article 4 in one main row]; article
5 ordinary in si/e. Right mandibular incisor with 5
teeth; molar not trilurative, small [Pbearing 4 or
more splayed, scmiarticulate spines, usually not
bearing fuzz]; palpar hump small. Palp of maxilla 1
biarticulate; inner plate with 2 setae. Setation of
maxilla 2 ordinary. Inner plate of maxillipeds ordi-
nary; apex of palp article 3 not protuberant, dactyl
elongate, apical nail distinct, elongate, bearing
large partner. Gnathopods ordinary, small, similar;
article 5 free, elongate, with weak eusirid attach-
ment, palms oblique to almost transverse, hand
ovatorectangular, heavily setose anteriorly but not
trichophoxid in shape. Article 2 of pereopod 3 of
broad form, articles 4-5 of pereopods 3-4 broad,
article 2 of pereopods $-4 not setose posteriorly;
pereopod 5 ordinary, article 2 naked ventrally, arti-
cle 3 slightly enlarged, dactyl normal. Epimera 1-2
lacking numerous long posterior setae, without
midfacial setae above ventral facial ridge; epimeron
3 ordinary. Urosomite 1 generally naked; urosomite
3 without dorsal hook or special process. Peduncle
of uropod 1 normally elongate, without apicoven-
tral spike, without special enlarged apicolateral or
medial spine; [Ppeduncular apices of uropods 1-2
not combed]; inner ramus of uropod 1 lacking mar-
ginal spines, some rami of uropods 1-2 continu-
ously spinose to apex, inner ramus of uropod 2
ordinary. Uropod 3 ordinary, elongate article 2 of
outer ramus carrying 2 long apical setae. Telson
elongate, with 2 apical spines or setae on each lobe
plus setules, with special lateral spines or setae.
DESCRIPTION.?Rostrum constricted, small to ob-
NUMBER 245 415
solescent. [Fuzz on article 1 of antenna 1 in male
unknown, calceoli on male antennae 1-2 unknown.
Prebuccal parts unknown. Right lacinia mobilis
unknown;] mandibular palp thin, article 1 short,
article 2 without outer setae, apex of article 3
oblique. [PLower lip bearing cones. Outer plate of
maxilla 1 unknown.] Inner plates of maxilliped
thick, ordinarily setose. Coxae 2-4 without special
anterodorsal hump. Some posterior spines on arti-
cle 6 of pereopods 1-2 thin and seta-like, [midapical
spine or seta unknown]. Article 2 of pereopod 5
without facial setae. Peduncle of uropod 1 with
dorsolateral spine confined apically, medial spine
confined apically, peduncle of uropod 2 with me-
dial spines widely spread. Peduncle of uropod 3
lacking extra subapical setae or spines. [PTelson
with ordinary pair of midlateral or dorsal setules
on each side unknown.]
TYPE-SPECIES.?Microphoxus minimus J. L.
Barnard, 1960 (monotypy).
COMPOSITION.?Unique.
RELATIONSHIP.?This genus differs from Birubius
in the shorter article 2 of antenna 1, the elongate
telson bearing lateral setae, the elongate article 2
on the outer ramus of uropod 3, the strong anterior
setation on the hands of the gnathopods, the reduc-
tion of setae on the inner plate of maxilla 1; the
large accessory nail on the dactyl of the maxilliped,
the slightly enlarged article 3 of pereopod 5 and the
widespread medial spination on the peduncle of
uropod 2. There may be distinctions in the lacinia
mobilis and molar. The type-species requires exten-
sive reanalysis.
From Parharpinia and Protophoxus, Micro-
phoxus differs in the absence of a special spine on
the peduncle of uropod 1, the broad articles 4-5 of
pereopods 3^1, the elongate telson and the strong
anterior setation on the hands of the gnathopods.
Microphoxus may have some relationship to the
Tipimegus group of genera in view of gnathopod 1,
pereopod 5 and the shape of pereopod 4, but
Microphoxus differs from those genera in the some-
what broader hands of the gnathopods, the absence
of posterior setae on article 2 of pereopod 4 and
epimera 1-2, the absence of facial setae on pereo-
pod 5, the absence of a spike on uropod 1, the short
article 2 of antenna 1, the strong nail of the maxil-
lipedal palp, the regular shape of article 2 on
pereopod 5, and the normal article 5 of antenna 2.
See Metharpinia below for further remarks.
Microphoxus minimus J. L. Barnard
Microphoxus minimus J. L. Barnard, 1960:291, pi. 46.
DISTRIBUTION.?Pacific Costa Rica, 4-9 m.
Metharpinia Schellenberg
Metharpinia Schellenberg, 1931:65.
DIAGNOSIS.?Eyes present. Flagella of antenna
1-2 unreduced in female. [Article 2 of antenna 1
unknown precisely, possibly ordinary], ventral setae
widely spread. [?Article 1 of antenna 2 not ensi-
form; article 3 with 2 setules; Pfacial spines on arti-
cle 4 in one main or 2 or more rows; article 5 ordi-
nary in size. Right mandibular incisor with 3 teeth;]
molar not triturative, small, elongate, conical,
[bearing unknown number of "numerous" spines,
Pusually not bearing fuzz]; palpar hump small.
Palp of maxilla 1 biarticulate; inner plate with 2
setae. Setation of maxilla 2 weak; inner plate of
maxillipeds ordinary; apex of palp article 3 not
protuberant, dactyl elongate, apical nail distinct,
immense. Gnathopods small, similar, but gnatho-
pod 2 weakly enlarged; article 5 of gnathopods 1-2
free, elongate, without eusirid attachment; palms
oblique, hands ordinary, ovatorectangular, moder-
ately setose anteriorly. Article 5 of pereopods 1-2
setose posteroproximally. Article 2 of pereopod 3
of broad form, articles 4-5 of pereopods 3-4 broad,
article 2 of pereopods 3-4 not setose posteriorly;
pereopod 5 ordinary, article 2 naked ventrally, arti-
cle 3 ordinary, dactyl normal. [PPeduncle of uropod
1 normally elongate, without apicoventral spike,
without special enlarged apicolateral?medial spine;
peduncular apices of uropods 1-2 not combed; inner
ramus of uropod 1 with marginal spines in one
row,] some rami continuously spinose to apex, in-
ner ramus of uropod 2 ordinary. Uropod 3 ordi-
nary, bearing elongate article 2 of outer ramus,
[Pcarrying 2 medium apical setae]. Telson elongate,
with 2+ apical spines or setae on each lobe plus
setules, with special lateral setae. [PEpimera 1-2
lacking long posterior setae, without midfacial setae
above ventral facial ridge]; epimeron 3 ordinary.
[PUrosomite 1 generally naked]; urosomite 3 without
dorsal hook or special process.
DESCRIPTION.?Rostrum fully developed, con-
stricted. [PFuzz on article 1 of antenna 1 in male
unknown; calceoli on male antennae 1-2 unknown.
416 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Prebuccal parts ordinary. PRight lacinia mobilis
bifid, thin;] mandibular palp thin, article 1 short,
article 2 without outer setae, apex of article 3
oblique. [PLower lip bearing cones. Outer plate of
maxilla 1 with 11 spines, one spine especially thick-
ened.] Inner plates of maxilliped thick, ordinarily
setose. Coxae 2-4 without special anterodorsal
humps. [PSome posterior spines on article 6 of
pereopods 1-2 thin and seta-like, midapical spine
or seta unknown.] Article 2 of pereopod 5 without
facial setae. [PPeduncle of uropod 1 with dorso-
lateral spines confined apically, medial spines con-
fined apically; peduncle of uropod 2 with only one
medial spine or setule confined apically. Peduncle
of uropod 3 lacking extra subapical setae or spines.
Telson with ordinary pair of midlateral or dorsal
setules on each side.]
TYPE-SPECIES.?Metharpinia longirostris Schel-
lenberg, 1931 (here selected on the basis of page
priority).
COMPOSITION.?Unique. Metharpinia cornuta
Schellenberg, 1931, is removed to a new genus to
be described (J. L. Barnard, in prep.).
REMARKS.?The diagnosis and description of the
genus are sprinkled liberally with brackets contain-
ing questioned guesswork about many morpholo-
gical characters not adequately described.
RELATIONSHIP.?Metharpinia and Microphoxus
appear to be closely similar genera in terms of tel-
son, maxillipedal dactyl, pereopods 3-4, article 3
of pereopod 5, molars, outer ramus of uropod 3,
gnathopods, inner plate of maxilla 1 and to some
extent the head.
Microphoxus differs only in the lack of posterior
setae on coxa 4 but so many characters of either
genus are unknown that further comments cannot
be made.
Metharpinia longirostris Schellenberg
Metharpinia longirostris Schellenberg. 1931:65-67. fig. 54;
1935:252.
Paraphoxus longirostris.?J. L. Barnard. 1960:275. pi. 43.
DISTRIBUTION.?South America, Valparaiso, Cor-
ral, and south of mouth of La Plata River, 9-13 m.
PHOXOCEPHALINAE
DIAGNOSIS.?Article 2 of antenna 1 usually short-
ened; mandibular molar fully triturative or re-
duced to a small hump with articulate spines; palp
of maxilla 1 uniarticulatc; solution on maxilla 2
reduced; gnathopod 2 significantly enlarged, or as
small as gnathopod 1; article 2 of pereopod 3 of
broad form; pereopod 5 ordinary.
DESCRIPTION.?Article 5 of antenna 2 usually re-
duced in size; epimeron 3 of rounded classification;
apices on peduncles of uropods 1-2 combed or not.
TYPE GENUS.?Phoxocephalus Stebbing.
COMPOSITION.?Diogodias, new genus; Hopi-
phoxus, new genus; Japara, new genus; Jerildaria,
new genus; Kondoleus, new genus; Leptophoxoides
J. L. Barnard; Leptophoxus Sars; Limnoporeia
Fearn-Wannan; Metaphoxoides Ledoyer; Meta-
phoxus Bonnier; Rikkarus, new genus; Uldanamia,
new genus; Vasco, new genus.
Key to the Genera of Phoxocephalinae
1. Article 3 of palp on maxilliped immensely produced 2
Article 3 of palp on maxilliped unproduced or scarcely so 5
2. Mandibular molar triturative Leptophoxoides
Mandibular molar not triturative Leptophoxus
3. Mandibular molar triturative 4
Mandibular molar not triturative, bearing spines, or molar absent 5
4. Hand-wrist of gnathopods eusirid, greatly enlarged, palpar hump of mandible huge, article
2 on outer ramus of uropod 3 very short Jerildaria, new genus
Hand-wrist of gnathopods weakly eusirid or not, not highly enlarged, palpar hump of man-
dible small to medium, article 2 on outer ramus of uropod 3 elongate Phoxocephalus
5. Rami of uropod 3 reduced, shorter than peduncle, outer ramus lacking article 2
Japara, new genus
Rami of uropod 3 ordinary 6
6. Flagellum of female antenna 2 reduced to one article, rostrum obsolete, article 2 of antenna
1 slightly elongate and produced apically, maxilliped with basal spouts, uropods 1-2 with
dorsal setae, inner ramus of uropod 2 reduced Kondoleus, new genus
NUMBER 245 417
Flagcllum of female antenna 2 multiarticulate, rostrum present, article 2 of antenna 1 short,
unproduced, maxilliped lacking basal spouts, uropods 1-2 with normal rami and lacking
dorsal setae 7
7. Gnathopods strongly chelate 8
Gnathopods subchelate, ocassionally gnathopod 1 weakly parachelate 9
8. Wrist of gnathopod 2 highly elongate, dactyls of pereopods 1-2 vestigial
Uldanamia, new genus
Wrist of gnathopod 2 not elongate, dactyls of pereopods 1-2 fully developed .. .limnoporeia
9. Article 2 on outer ramus of uropod 3 vestigial Metaphoxoides
Article 2 on outer ramus of uropod 3 well developed 10
10. Epimeron 3 grossly serrate, hand of gnathopod 2 thin, sinuous, palm extremely oblique
Rikkarus, new genus
Epimeron 3 of rounded classification, hand of gnathopod 2 expanded, palm weakly to mod-
erately oblique 11
11. Rami of uropods 1-2 dorsally setose near apices Hopiphoxus, new genus
Rami of uropods 1-2 neither setose nor bearing dorsal armaments near apices 12
12. Article 5 of gnathopods elongate Diogodias, new genus
Article 5 of gnathopods short 13
13. Dactyls of pereopods 1-2 stunted Vasco, new genus
Dactyls of pereopods 1-2 elongate Metaphoxus
Phoxocephalus Stebbing
Phoxus Kr0yer, 1842:150 [not Billberg 1820, Coleoptera].
Phoxocephalus .Stebbing, 1888:810-811 [new name].
DIAGNOSIS.?Eyes present but occasionally defec-
tive. Flagellum of antenna 2 reduced in female.
Article 2 of antenna 1 ordinary to shortened,
ventral setae confined apically. Article 1 of antenna
2 not ensiform; article 3 with 2 setules; facial
spines on article 4 in 2 or more rows, plus special
apical spines; article 5 thin and short. Right man-
dibular incisor with 2-3 teeth; molar strongly tri-
turative, large to medium; palpar hump small to
medium. Palp of maxilla 1 uniarticulate; inner
plate naked. Setation of maxilla 2 weak. Inner
plate of maxillipeds ordinary; apex of palp article
3 not protuberant, dactyl elongate to medium,
apical nail distinct, short to elongate. Gnathopods
dissimilar, gnathopod 2 weakly to strongly en-
larged, article 5 of gnathopod 1 of ordinary length,
very short and cryptic on gnathopod 2, with weak
eusirid attachment; palms oblique to transverse,
hands of gnathopods 1-2 ordinary, ovatorectang-
ular to broadened, poorly setose anteriorly. Article
5 of pereopods 1-2 setose posteroproximally. Arti-
cle 2 of pereopod 3 of broad form, articles 4-5 of
pereopods $-A narrow to medium, article 2 of
pereopods 3-4 not setose posteriorly; pereopod 5
ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Epimera 1-2 lacking long pos-
terior setae, without midfacial setae above ventral
facial ridge; epimeron 3 of rounded classification
and bearing 3 or fewer long setae. Urosomite 1
generally naked; urosomite 3 without dorsal hook
or special process. Peduncle of uropod 1 normally
elongate, without apicoventral spike, without spe-
cial enlarged apicolateral or medial spine, pedun-
cular apices of uropods 1-2 combed; inner ramus
of uropod 1 with marginal spines in one row, no
rami continously spinose to apex, apical nails
poorly developed, inner ramus of uropod 2 ordi-
nary. Uropod 3 ordinary, elongate article 2 of outer
ramus carrying 1-2 short apical setae. Telson
elongate, with 1-2 apical spines or setae on each
lobe plus setules, rarely with special dorsal and
lateral spines or setae.
DESCRIPTION.?Rostrum fully developed. Fuzz on
article 1 of antenna 1 in male present; calceoli on
male primary flagellum of antenna 1 present.
Calceoli on article 5 of male antenna 2 present,
flagellum in male with calceoli. Prebuccal parts
ordinary, poorly separated from each other, upper
lip dominant. Right lacinia mobilis bifid, flabel-
late; mandibular palp medium to thin, article 1
short, article 2 without outer setae, apex of article
3 oblique. Lower lip lacking cones. Outer plate of
maxilla 1 with 7-9 spines, no spine especially thick-
ened. Inner plates of maxilliped especially thin,
poorly armed. Coxae 2-4 without special antero-
dorsal humps. Some posterior spines on article 6 of
pereopods 1-2 thin and seta-like, others thick and
stiff, midapical spine or seta present. Article 2 of
418 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pereopod 5 without facial setae. Peduncle of
uropod 1 with dorsolateral spines confined apically,
medial spines confined apically or widely spread;
peduncle of uropod 2 with only one medial spine
or setule confined apically or with spines widely
spread. Peduncle of uropod 3 lacking extra sub-
apical setae or spines. Telson with ordinary pair of
midlateral or dorsal setules on each side.
TYPE-SPECIES.?Phoxus holbolli Kr0yer 1842
[selected by Boeck, 1876].
COMPOSITION.?Phoxus bassi Stebbing, 1888;
Phoxocephalus burleus, new species; Phoxoceph-
alus homilis J. L. Barnard, 1960; Phoxocephalus
keppeli, new species; Phoxus kergueleni Stebbing,
1888; Phoxocephalus kukathus, new species:
Phoxocephalus regium K. H. Barnard, 19S0;
Phoxocephalus rupullus, new species; Phoxoceph'
aim tenuipes Stephensen, 1925; Phoxocephalus
tunggeus, new species.
REMARKS.?The type-species has such small
gnathopods, scarcely divergent in size, that the
other known species of the genus might be re-
moved to a distinctive subgenus at least.
A diversity of dactyls on the maxilliped is alto
present and the medial spination on the peduncles
of uropods 1-2 is variable.
Key to the World Specie* of Phoxocephalus
(Not including P. keppeli)
1. Hand of gnathopod 2 only 1.1 time* as long as wide 2
Hand of gnathopod 2 at least 1.7 times as long as wide S
2. Eyes present, hand of gnathopod 1 about 1.9 times as long as wide P- bmad
Eyes absent, hand of gnathopod 1 about 1.2 times as lung as wide P. tenuipet
3. Eyes absent or structurally deficient, gnathopods small, gnathopod 2 scarcely larger than
gnathopod 1 P. holboUi
Eyes present, large, gnathopod 2 enlarged 4
4. Outer ram us of uropod 1 as long as inner ramus and dorsally spinose 5
Outer ramus of uropod 1 shortened and naked 7
5. Facial setae of epimeron 2 confined to anterior third of surface P. regutm
Facial setae of epimeron 2 spread fully posterior 6
6. Cuticle with villi organized into polygons P. rupullut, new species
Cuticle with unpatterned striations P. burtew, new species
7. Epimeron 3 with long posterior setae, apical setae on article 2 of outer ramus on uropod S
elongate P. tunggeut, new species
Epimeron 3 with short posterior setules or none, apical setae on ankle 2 of outer ramus on
uropod 3 very short 8
8. Outer ramus of uropod 2 with 3 setae P. homilis
Outer ramus of uropod 2 naked 9
9. Inner ramus of uropod 1 with 3 dorsal spines P. ktrgueletd
Inner ramus of uropod 1 with one dorsal spine P. kukathus, new species
Phoxocephalus bassi (Stebbing)
FICURES 213, 214
Phoxus bassi Stebbing, 1888:811-815, pi 54.
Phoxocephalus bassi.?Stebbing, 1906:135; 1910:577, 635.?
J. L. Barnard, 1960:298.
Not Phoxocephalus bassi.?K. H. Barnard, 1930:330-331, fig.
9b [see P. keppeli, new species].
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about 55
percent of length, rostrum reaching apex of article
3 on antenna 1, unconstricted, tapering almost to
point, apex rounded. Eyes of medium size, clear of
pigment core in preservative. Article 1 on peduncle
of antenna 1 almost 1.9 times as long as wide, 1.8
times as wide as article 2, apicoventral margin with
several setules, strongly produced dorsal apex with
3 setules; article 2 about 0.45 times as long as arti-
cle 1, apicoventral surface with 5 setae, article 3
attached almost terminally to article 2; accessory
flagellum slightly shorter than primary flagellum,
5-articulate, primary flagellum ^-articulate, aesthe-
tascs absent. Spine formula on article 4 of antenna
2 = 1-3-4-5, proximal sets with one accessory
setule, spines thin, dorsal margin with set of 3
setae, long to short, ventral margin with 5 subbasal
penicillate setules and about 17 long or short setae,
NUMBER 245 419
FIGURE 213.?Phoxocephalus bassi (Stebbing), female "a," 9.5 mm.
420 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 214.?Phoxocephalus bassi (Stebbing), female "a," 9.5 mm.
NUMBER 245 421
apex with 2 short spines; article 5 about 0.7 times
as long as article 4, facial spine formula = 2, with
one accessory setule, full distal margin with 4 short
to long spines, dorsal margin smooth, ventral
margin with 4 sets of 1-2 long and short setae;
fiagellum scarcely longer than article 4 of peduncle,
6-articulate. Palpar hump of mandible large; right
incisor bidentate, distolateral tooth obsolete; left
incisor bidentate, distal branch weakly trifid; right
lacinia mobilis Habellate, distal branch with
toothed margin, face with humps; left lacinia
mobilis flabellate, trifid; right rakers 2, left 3;
molar triturative, on medium-length pedicle; palp
article 2 with 3 subbasal setae and 2 apical setae on
inner margin, article 3 about 0.9 times as long as
article 2, oblique apex with 11 setae. Mandibular
lobes on lower lip of medium length, weakly at-
tenuate and apically rounded, inner lobes fused
together for most of their length but with distinct
raphe and turgid faces accentuating raphe. Inner
plate of maxilla 1 enlarged, elliptical, naked; outer
plate with 9 spines; palp scarcely exceeding outer
plate, bearing 2 apical, one apicolateral, and 2
apicomedial setae, some setae strap-shaped. Inner
plate of maxilla 2 much broader than and weakly
exceeding outer plate, medial margin of inner plate
with about 8-10 short setae, outer plate with 8
apical and medial setae and 2 apicolateral setae.
Inner plate of maxilliped with one medial spine, 2
stout and 3 slender apical spines; outer plate with
6 stout and one slender medial spines, one apico-
lateral seta; article 2 of palp moderately setose
medially, article 3 with 4 facial setae, article 4 of
medium length, apical spine short, bearing 2 ac-
cessory setules. Coxa 1 distally expanded and
broadly rounded anteroventrally; long setal for-
mula of coxae 1-4 = 12-11-10-6; anterior and pos-
terior margins of coxa 4 weakly divergent, poster-
ior margin weakly convex, posterodorsal corner
subsharp, posterodorsal excavation of medium size
and L-shaped, ratio of width to length = 19:21.
Gnathopod 2 much stouter than gnathopod 1; long
posterior setae on article 2 of gnathopods 1-2 = 3
and 7, short posterior setae = 1 and 0, short anter-
ior setae = 0 and (3-4); article 3 weakly elongate
on gnathopod 1, gnathopods subeusirid; article 5
thin; article 6 attached to produced apex of article
5, article 6 of gnathopod 1 elongate and rectang-
ular, of gnathopod 2 trapezoidal, nearly as broad
as long [broader than long if length measured as
true longitudinal facial axis]; palm oblique, defin-
ing hump large on gnathopod 2, small on gnatho-
pod 1, dactyl fitting palm, apex of dactyl on
gnathopod 1 with flake [but apex of dactyl on
gnathopod 2 eroded on unique specimen]; poster-
ior margin of article 5 free on gnathopod 1; length
ratios of articles 5-6 on gnathopods 1-2 = 54:64
and 46:93 [midfacial axis =46:78], width ratios
= 22:38 and 21:83. Pereopods 1-2 similar to each
other; long posterior setae on article 2 of pereopods
1-2 = 4 and 5, short posterior setae = 1 and 2,
short anterior setae = 2 and 2; article 4 with 4 and
5 facial setae, posterior margins of article 4 mod-
erately setose; article 5 with long distal spine reach-
ing to M.85 on article 6, with very small contiguous
spine at base, article 5 with 3 and 3 facial setae on
pereopods 1-2; article 6 of pereopods 1-2 with
rows of 5 and 2 or 5 and 3 spines plus spine mid-
distally, spines mostly long; dactyl with apical nail
distinct and deeply immersed, inner margin with
weak acclivity and short setule. Coxae 5-7 postero-
ventral setule formula = 1-1-3. Width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 53:22:21:10, of
pereopod 4 = 69:26:19:10, of pereopod 5 = 83:
17:16:9, length ratios of articles 2, 4, 5, 6 of
pereopod 3 = 67:21:27:33, of pereopod 4 = 84:
55:48:59, of pereopod 5 = 96:18:28:24; article 2
of pereopod 5 extending to apex of article 4;
apex of article 6 finely combed medially. Epi-
mera 1-2 with facial ridge, each with rounded-
quadrate posteroventral corner and weakly convex
posterior margin; epimeron 3 with setule notch at
posteroventral corner, posterior margin broadly
rounded, bearing 3 setae in notches; epimeron 1
with 2 posteroventral setae in horizontal tandem
(marked by dots in illustration); epimeron 2 with
horizontal crescent of 6 setae. Urosomite 1 bearing
one seta at base of uropod 1, urosomites almost
flush dorsally, urosomite 1 with one seta on each
side dorsally. Peduncle of uropod 1 with one small
basofacial seta, lateral apex with short spine, med-
ial margin with 4 spine-setae, distalmost elongate;
rami of uropods 1-2 with articulate immersed
apical nail and complex flake, inner ramus of uro-
pod 2 with one dorsal spine, all other rami with 2
dorsal spines, inner ramus of uropod 1 with sub-
basal setule (marked by dot on illustration), outer
ramus of uropod 1 reaching farther than inner
ramus; peduncle of uropod 2 with 4 dorsal spines,
apicomedially with one spine, apex of peduncle on
422 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
uropod 2 with comb, absent on uropod 1. Uropod
3 long, rami feminine, peduncle with 5 ventro-
lateral spines, medially with 2 apical setules, inner
ramus extending almost to M. 70 on article 1 of
outer ramus, apex with ?2 setae [broken off], article
2 of outer ramus weakly elongate, 0.28, bearing 2
short apical setae, outer margin of article 1 with 4
acclivities, spine formula = 1-1-1-1-2, medial apex
with spine. Telson elongate, length-width ratio
14:11, apices weakly tapering, with weak apico-
medial lobe, lateral acclivity bearing long spine
and shorter lateral setule, main spine in middle
of apex, dorsal pair of diverse plusetules near M.
33, additional pair of long lateral setae on each
side near M. 70. Bifid bulbar setules of cuticle ex-
tremely sparse.
DESCRIPTION OF MALE.?See Stebbing (1888).
OBSERVATIONS.?Prebuccal complex weakly con-
vex anteriorly, weakly extended forward; on
unique female in hand, apicoventral spine on inner
plate of maxilliped occuring between either main
spines 1-2 or 2-3, differing on the two plates.
ILLUSTRATIONS.?Specimen with several defects:
eye sunken, full lateral dimensions not shown;
lower lip damaged, outline of outer lobe lacking
armaments; palp and outer plate of maxilla 1 dam-
aged, illustration of restored view.
VOUCHER MATERIAL.?AM P. 18243, female "a,"
9.5 mm (illus.), off Sydney, New South Wales, Aus-
tralia, 18 Jun 1962, 150 m.
REMARKS.?Stebbing (1888) described the male of
this species, about 10 mm long, from Bass Strait.
Our female apparently belongs to P. bassi, fitting
the a priori concept of a female of the species in
dozens of significant characters. Stebbing shows the
male uropod 2 bearing more than twice as many
spines on the peduncle of uropod 2 than in our
female, shows 3 spines on the outer ramus of uro-
pod 2 and none on the inner. Stebbing's view of
uropod 1 is apparently lateral whereas his view of
uropod 2 is apparently medial; 2 ventral pedunc-
ular spines on uropod 2 are interpreted by us to
be medial peduncular spines; we visualize the
peduncle as heavily flattened in Stebbing's view,
forcing the spines ventrad.
Differences in spine counts on uropods 1-2: are
common distinctions between males and females of
various phoxocephalids.
The mandibles of this species, as seen in our
specimen and that of Stebbing, are heavily cal-
cified, with the proximal tooth on the incisors of
both sides being reduced in prominence.
K. H. Barnard's identification of P. bassi from
New Zealand is removed to a new species, P. kep-
peli (see below).
MATERIAL.?AM, one sample (1).
DISTRIBUTION.?Southeastern Australia, Bass
Strait, Wata Mooli; New South Wales, off Sydney;
surface to 150 m.
Phoxocephalus kukathus, new species
Fict u s 215-217
DESCRIPTION OF FEMALE.?Head about 19 per-
cent of total body length, greatest width about 55
percent of length, rostrum reaching middle of arti-
cle 2 on antenna 1, unconstricted, weakly tapering,
apically rounded. Eyes large, red in preservative,
clear of pigment core. Article 1 on peduncle of
antenna 1 about 1.5 times as long as wide, twice as
wide as article 2, apicoventral margin with several
setules, strongly produced dorsal apex with 4
setules; article 2 about 0.55 times as long as article
1, apicoventral surface with 4 setae, article 3 at-
tached to lateral face of article 2 with override of
article 2 medially; accessory flagellum shorter than
primary flagellum, 3-articulate; primary flagellum
4-articulate, aesthetascs absent. Spine formula on
article 4 of antenna 2 = 1-3-5-3 or 1-3-5-4 or
1-3-4-2, proximal set with 1-2 accessory setules,
next set with one accessory setule, spines thin,
dorsal margin with one spinule, ventral margin with
2 subbasal penicillate setules and 7-9 long or short
setae, apex with 3 thin, short spines; article 5 less
than 0.6 times as long as article 4, facial spine
formula = 3 or 2, with one accessory setule, dorso-
distal corner with 3 spines, dorsal margin smooth,
ventral margin with 2 pairs of long-short setae,
ventrodistal apex with one long, 2 short setae;
flagellum shorter than article 4 of peduncle, 4-artic-
ulate. Pal par hump of mandible small; right in-
cisor weakly tridentate, with weak midnotch, acces-
sory notch near tooth 2, left basically bidentate,
distal branch weakly quadridentate and bearing
weak proximal facial humps; right lacinia mobilis
flabellate, distal branch poorly defined, margin
toothed, face with or without weak hump; left
lacinia mobilis flabellate, irregularly 4-5 toothed;
right rakers 2, left 3; molar triturative, on long
NUMBER 245 423
s
FIGURE 215.?Phoxocephalus kukathus, new species, male "c," 4.76 mm (u ? holotype, female
"u," 4.40 mm).
424
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 216.?Phoxocephalus kukathus, new species, male "c," 4.76 mm (u ? holotype, female
"u," 4.40 mm).
NUMBER 245 425
r
FIGURE 217.?Phoxocephalus kukathvs, new species, male "c," 4.76 mm (u ? holotype, female
"u," 4.40 mm).
pedicle; palp article 2 with one small subbasal seta
and 3 subapical setae on inner margin, article 3
about 0.85 times as long as article 2, oblique apex
with 10-11 setae. Mandibular lobes on lower lip of
medium length, weakly attenuate and apically
rounded, inner lobes fused together for most of
their length but with distinct raphe. Inner plate of
maxilla 1 large, elliptical, naked; outer plate with
426 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
7 spines; palp exceeding apex of outer plate, bear-
ing 3 apical and one apicolateral strap-shaped setae.
Inner plate of maxilla 2 much broader and slightly
shorter than outer plate, medial margin of inner
plate with one disjunct and distinctly medial seta
plus 4-5 other short setae on apicomedial margin,
perhaps truly apical setae shifted medially; outer
plate with 3-5 apical and medial setae. Inner plate
of maxilliped with one mediofacial spine and 2
apical spines; outer plate with 4 spines; article 2 of
palp with sparse and stout medial setae, article 3
with one facial seta, article 4 long, apical spine
short, bearing 2 accessory setules. Coxa 1 distally
expanded and broadly rounded anteroventrally;
long setal formula of coxae 1-4 = (5-8)-(5-8)-
(5-7)-(2-5); anterior and posterior margins of coxa
4 almost parallel, posterior margin weakly convex,
posterodorsal corner rounded, posterodorsal excava-
tion of medium size and L-shaped, ratio of width
to length = 16:17. Gnathopod 2 much stouter than
gnathopod 1; long posterior setae on article 2 of
gnathopods 1-2 == 5 and 5; gnathopod 1 subeusirid,
article 3 weakly elongate, article 5 thin, elongate,
article 6 attached to produced apex of article 5,
article 6 stout, ovatorectangular, palm oblique, de-
fining hump large on gnathopod 2, small on
gnathopod 1, dactyl weakly overlapping palm,
apices of dactyls with flake, posterior margin of
article 5 on gnathopod 1 almost cryptic; length
ratios of articles 5-6 on gnathopods 1-2 = 46:69
and 39:101, width ratios = 19:47 and 19:66. Pereo-
pods 1-2 similar to each other; long posterior
setae on article 2 of pereopods 1-2 = 4-6 and 4-7;
article 4 with about 6 apicoanterior setae, 2 of these
facial, posterior margins of articles 4-5 moderately
setose; article 5 lacking facial setae, with long distal
spine reaching to M. 100 on article 6, with very
small contiguous spine at base; article 6 with one
long posterior setal-spine and rows of 3 and 2 pos-
terodistal setal-spines, some sharp, others rounded
apically, dactyl with apical flake and thin inner
setule appressed in marginal groove (pulled out to
illustrate), apical nail almost fully fused to dactyl.
Coxae 5-7 .posteroventral setule formula - 1-1-1.
Width ratios of articles 2,4, 5, 6 of pereopod 3 =
60:18:15:8, of pereopod 4 = 66:18:12:6, of pereo-
pod 5 = 94:17:14:8, length ratios of pereopod
3 = 70:27:22:25, of pereopod 4 = 91:48:40:39, of
pereopod 5 = 116:25:27:16; article 2 of pereopod
5 extending to or exceeding apex of article 4;
apex of article 6 finely combed. Gill vestigial
or absent on pereopod 5. Epimera 1-2 with weak
facial ridge (omitted on drawing of female on epi-
meron 1), each with rounded-quadrate postero-
ventral corner and weakly convex posterior margin;
epimeron 3 with weak posteroventral tooth and
sinus bearing setule, posterior margin broadly
convex, bearing 2-3 setule notches; epimeron 2
with 2-4 facial setae in horizontal row. Urosomitc
1 bearing one lateral setule at base of uropod
1 and 1-3 proximoventral setae on each side,
urosomite 1 weakly produced dorsally and bear-
ing 2 setules; urosomite 3 with long seta at ven-
tral base of uropod 3. Peduncle of uropod I oc-
casionally with one basofacial seta, usually naked,
each apex of peduncle with one medium spine, no
medial marginal spines, inner ramus of urojxxl 1
and both rami of uropod 2 with apical flake and
articulate spine, rami of uropod 2 otherwise naked,
inner ram us of uropod I with one dorsal spine and
one mediobasal spinule, outer lamus of uropod 1
slightly shortened, thinner and more curved than
inner ramus, lacking spines, each apex of peduncle
on uropod 2 with medium spine, apex of peduncle
on uropods 1-2 with lateral comb. Uropod 3 long,
rami feminine, peduncle with 5 long ventrolateral
spines, medially with one short spine, inner ramus
extending to M. 70 on article 1 of outer ramus,
apex with 2 setae, article 2 of outer ramus elongate,
0.60, bearing one apical setule, outer margin of arti-
cle 1 with one spine-seta, apex with 2 spine-setae,
medial apex with one long spine-seta. Telson long,
length-width ratio about 6:5, apices tapering but
broadly truncate, weakly excavate, each bearing
long mid spine and small apicolateral setule, dorsal
pair of plusetules weakly diverse. Bulbar setules of
cuticle extremely sparse.
DESCRIPTION OF MALE.?Eyes much larger than
those of female. Article 1 of antenna 1 with medial
patch of fuzz; disparity in length of flagella greater
than in female, accessory flagellum 3-articulate,
primary flagellum 5-articulate, bearing several
aesthestascs, one calceolus (6-chambered) each on
articles 1-2 and highly elongate apical seta on arti-
cle 5. Spine formula on article 4 of antenna 2
peduncle = 4-5-2 or 3-5-2, article 5 with 2-3 facial
spines and 2-3 apicodorsal spines, articles 3-4 with
dorsomedial fuzz, article 5 with one dorsal calceolus
and 2 groups of stiff male setae, ventrodistal margin
poorly setose; flagellar formula = (25-27), 1-4,6, 8
NUMBER 245 427
. . . penultimate, or 1-3, 5, 7 Article 2 of
mandibular palp with 3-6 long subapical setae.
Inner plate of maxilla 2 narrower than in female,
setae of both plates larger, setae more numerous on
outer plate. Mediofacial spine on inner plate of
maxilliped very rarely more distally placed than in
female. Long posterior setae on article 2 of gnatho-
pods 1-2 and pereopods 1-2 = (l-4)-(l-3)-{2-4)-
(2-4). Coxa 4 narrower than in female. Article 2 of
pereopxis 3-5 more slender than in female. Gill
well developed on pereopod 5. Epimera enlarged,
epimeron 2 with 2-4 facial setae, epimeron 3 with
2-4 posterior setule notches besides one at ventral
corner. Urosomc smaller than in female relative to
epimera, urosomite 1 with 2-5 proximoventral setae
on each side. Body generally longer than in female.
Uropod 1 with 1-2 basofacial setae. Raini of uropod
3 masculine, inner extending to M. 87 on article 1
of outer ramus, both margins setose, outer ramus
with setose medial margin on article 1, lateral
margin with pair of setae closely contiguous, article
2 with 2 apical setules. Telson slightly more elon-
gate than in female, with basal patches of dorsal
denticles.
OBSERVATIONS.?Additional variations in facial
spine formulas on article 4 of female antenna 2 =
1-2-5-2, 1-3-4-1, 1-3-1-3, and 1-2-1-2. Outer
plate of maxilliped on female, 4.1 mm, CPBS 33S,
with one outer plate with only 2 thick apical spines
and one thinner medial spine. Long posterior setae
on article 2 of female gnathopod 1 most commonly
4, but variable between 1-5; on gnathopod 2 vary-
ing between 2 and 5. Epimeron 3 of female excep-
tionally with either 0 or 4 posterior notches besides
posteroventral notch. Six known Western Port
males and SBS males of comparable size (4.5 mm)
differing weakly in shapes of gnathopods, Western
Port nek tonic male with long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
2-1-2-2, on SBS male = 4-2-3-4, such setae gen-
erally fewer in males than in females of comparable
size; hands of gnathopods 1-2 in Port Phillip males
and females generally thinner than those of West-
ern Port, Port Phillip specimens generally with
8-9 apical setae on mandibular palp compared with
10-11 on Western Port and SBS specimens.
ILLUSTRATIONS.?Apex on outer ramus of uropod
2 as illustrated for male probably aberrant, normal
condition as shown in other illustration.
HOLOTYPE.?NMV, female "u," 4.40 mm.
TYPE-LOCALITY.?CPBS 12N/3, 16 Mar 1965,
Western Port, Victoria, Australia, 3 m, Zostera,
mud.
VOUCHER MATERIAL.?Type-locality, female "b,"
3.77 mm; CPBS 12N/2, male "c," 4.76 mm (illus.).
RELATIONSHIP.?This species differs from P. bassi
(Stebbing, 1888), a sympatriot from Bass Strait, in
the longer article 6 of gnathopod 2 which in P.
bassi is as broad as long, in the absence of an
oblique row of facial setae on epimeron 3, and in
the slightly shortened and thin outer ramus of
uropod I.
Phoxocephalus kukathus is strongly allied to P.
kergucleni (Stebbing, 1888), from Kerguelen Island,
and may not be specifically distinct from that
species. Only one strong distinction can be dis-
cerned in that species, the presence of 3 dorsal
spines on the inner ramus of uropod 1, in contrast
to one spine in P. kukathus. The right mandibular
incisor of P. kergueleni is more strongly toothed
and notched than that of P. kukathus. There may
be distinctions in the breadth of plates on maxilla
2 and distinctions in uropod 3 but the "female"
illustrated by Stebbing has a uropod 3 character-
istic of so-called intersexes and may therefore be
senile or aberrant; the plates of maxilla 2 in Steb-
bing's illustrations are closer to the male depicted
herein than to the female. Stebbing has either over-
looked setae on epimeron 2 or else they are absent
in P. kergueleni, a character of good specific value
which should be verified.
Phoxocephalus regium K. H. Barnard, 1930, from
New Zealand, has the rami of uropod 1 extending
equally.
MATERIAL.?CPBS, 15 samples from 9 stations
(53); WPBES, 6 samples from 4 stations (10); RHM
1 sample (1); PPBES, 48 samples frqm 24 stations
(249); SBS, 13 samples from 9 stations (132); AM,
one sample (14).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, intertidal to 22 m, sand, silty sand,
sandy silt, clayey silt, clay. New South Wales, off
Malabar, 48-75 m, sand, sandy mud, sandy gravel,
South Australia, Kangaroo Island, neritic.
Phoxocephalus tunggeus, new species
FIGURES 218, 219
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width about 55
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s
FIGURE 218.?Phoxocephalus tunggeus, new species, holotype, female "a," 5.66 mm.
NUMBER 245 429
Rl
FIGURE 219.?Phoxocephalus tunggeus, new species, holotype, female "a," 5.66 mm.
430 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
percent of length, rostrum reaching apex of article
2 on antenna 1; unconstricted, tapering evenly and
strongly, apex rounded, very narrow. Eyes of me-
dium size, clear of pigment core. Article 1 on pe-
duncle of antenna 1 almost 1.4 times as long as
wide, 2.2 times as wide as article 2, apicoventral
margin with several setules, strongly produced
dorsal apex with 3 setules; article 2 about 0.42
times as long as article 1, apicoventral surface with
3 setae; article 3 attached terminally to article 2;
accessory flagellum much shorter than primary fla-
gellum, 3-articulate; primary flagellum 5-articulate,
aesthetascs present. Spine formula on article 4 of
antenna 2 = 3-5-5-1, middle sets each with one
accessory setule, proximal set with 3 accessory set-
ules, spines thin, dorsal margin with 2 sets of 1-3
setae, ventral margin with about 9 sets of 1-2 setae
and 3 subbasal penicillate setules, apex with one
thin spine; article 5 slightly more than 0.5 times as
long as article 4, facial spine formula = 3, with one
accessory setule, dorsodistal corner with 2 setae and
3 spines, apex with one spine, ventral margin with
5 sets of 1-2 setae, ventrodistal apex with one long
seta; flagellum about 0.85 times as long as article
4 of peduncle, 4-articulate. Palpar hump of mandi-
ble small-medium; right incisor tridentate, with 2
notches between teeth 1 and 2; left basically biden-
tate, distal branch weakly 4-dentate; right lacinia
mobilis flabellate, distal branch poorly defined,
margin toothed, face with ragged hump-line; left
lacinia mobilis flabellate, 4-dentate; right rakers 2,
left 3; molar triturative, on long pedicle; palp arti-
cle 2 with 5 long setae on apical margin distally,
article 3 about 0.85 times as long as article 2,
oblique apex with 14 setae. Mandibular lobes of
lower lip of medium length, apically rounded, in-
ner lobes fused together for most of their length
but with distinct raphe. Inner plate of maxilla 1
large, elliptical, naked; outer plate with 7 spines;
palp exceeding apex of outer plate, bearing 3 apical
and 2 apicolateral setae. Inner plate of maxilla 2
almost twice as wide as outer plate, slightly exceed-
ing apex of outer plate, setae all medial; outer plate
with one apicolateral, 4 apical and 2 medial setae.
Inner plate of maxilliped with one mediofacial
seta, 3 apical spines; outer plate with 5 spines; arti-
cle 2 of palp with medium dense medial setae, arti-
cle 3 with 2 facial setae, dactyl long, apical spine
short, bearing 2 accessory setules. Coxa 1 distally
expanded, rounded anteroventrally; long setal
formula of coxae 1-4 = 5-7-6-6; anterior and pos-
terior margins of coxa 4 divergent, posterior mar-
gin oblique, convex, posterodorsal corner subsharp,
posterodorsal excavation of medium size and lun-
ate, ratio of width to length = about 11. Gnatho-
pod 2 much stouter than gnathopod 1; long poste-
rior setae on article 2 of gnathopods 1-2 ? 11 and
11-14, short posterior setae = 1-1, short ameriors =
3 and 5, long anterolaterals = 2 and 0, article S
short; only gnathopod 2 weakly eusirid, article 6
broadened but elongate, rectangular, palm oblique,
defining hump large only on gnathopod 2, dactyl
fitting palm (slight override), apices of dactyls with
flake, posterior margin of article 5 on gnathopod 1
free, length ratios of articles 5-6 on gnathopods
1_2 - 44:65 and 38:97. width ratios - 2241 and
22:73. Pereopods 1-2 alike, long posterior setae on
article 2 of pereopods 1-2 10-11 ami 13, short
anteriors 2 and 1, article 1 of |>ereo|>ods 1-2 with
4 apicolateral setae each. |>osterior margins of arti-
cle 4 moderately setose, of article 5 more densely
so, article 5 with 2 and 2 anterofacial setae, with
long posterodistal spine reaching to M. 110 on
article 6, with small contiguous seta-spine at base,
article 6 with one long posterior seta-spine, one
short and stouter middistal spine and rows of 2 and
2 long, thin apical spines on each of pereopods 1-2,
dactyl with long hooked apical flake bearing apical
setule, inner margin with subfacial seta emerging
into channel forming one limb of flake, apical nail
almost fully fused to dactyl. Coxae 5-7 postero-
ventral setule formula = 1-1-1. Width ratios of
articles 2, 4, 5, 6 of pereopod 3 =49:18:14:8, of
pereopod 4 = 57:24:14:11, of pereopod 5=89:17:
14:7, length ratios of pereopod 3 = 65:22:31:34,
of pereopod 4 = 92:50:73:unknown, of pereopod
5= 122=24:26:21; article 2 of pereopod 5 slightly
exceeding middle of article 5; apex of article 6
finely combed. Only epimeron 1 with facial ridge,
posteroventral corner rounded, posterior margin
convex, epimera 2-3 with rounded-quadrate pos-
teroventral corners, epimeron 3 with strongly con-
vex posterior margin bearing 3 setae on dorsal
half, epimeron 2 with weakly convex posterior
margin, epimera 1-2 each with midposterior setule,
epimeron 1 naked, epimeron 2 with one facial
anteroventral seta, ventral margin of epimeron
3 naked. Urosomite 1 bearing one lateral seta
at base of uropod 1, urosomite 2 with one such
seta, urosomites dor sally truncate, ventral base
NUMBER 245 431
of urosomite 3 with 3 long setae. Peduncle of uro-
pod 1 with 1-2 facial setae at M. 30-35, lateral apex
of peduncle with one spine, medial margin with 4
short spines, outer ramus short, reaching only M.
70 on inner ramus, curved, naked, apical nail
weakly denned, lacking flake (apparently apex
poorly preserved), inner ramus with 4 dorsal spines
and one mediobasal setule, apical nail articulate
but immersed and solidly fixed, bearing complex
flake; peduncle of uropod 2 with short apicolateral
spine and lateral margin with 4 long setae, [outer
ramus apically broken, occurrence of spines un-
known], inner ramus naked, apical nail as on
uropod 1 inner ramus; peduncular apices of uro-
pods 1-2 uncombed. Uropod 3 long, rami feminine,
peduncle with 3 long ventrolateral setae, dorsolat-
erally and dorsomedially with setule, inner ramus
extending to M. 70 on article 1 of outer ramus,
apex with one seta, article 2 of outer ramus elon-
gate, 0.44, bearing 2 apical setae, outer margin of
article 1 with 3 acclivities, elongate spine-seta
formula- 1-1-1-1, medial apex with one spine-
seta. Telson slightly elongate, length-width ratio
= 17:15, apices scarcely tapering, each with weak
medial protrusion and weak lateral acclivity bear-
ing one long spine-seta and one ordinary lateral
setule, dorsal pair of plusetules alike. Cuticle glass
smooth, very sparsely armed with ordinary bulbar
setules.
OBSERVATIONS.?Prebuccal complex, epistome and
upper lip poorly preserved, apparently like those
of P. kukathus, Figure 215: uB and uU.
ILLUSTRATIONS.?Outer ramus of uropod 2 broken
distally on both sides; apices of both fourth pereo-
pods damaged or missing.
HOLOTYPE.?AM, female "a," 5.66 mm. Unique.
TYPE-LOCALITY.?AM P. 18269, 20 Jun 1962, SW
of Cape Everard, Victoria, Australia, 38? 15' S,
149? 12' E, 300 m.
RELATIONSHIP.?Phoxocephalus tunggeus, being
based on a single individual, might appear to repre-
sent a senile condition of P. kukathus, new species,
but the following characters appear to distinguish
P. tunggeus adequately: (1) longer setae of epi-
meron 3; (2) absence of ridge and presence of only
one facial seta on epimeron 2; (3) presence of long
setae on peduncle of uropod 2; (4) complex scale on
dactyls of pereopods 1-2.
Phoxocephalus tunggeus differs from Californian
P. homilis J. L. Barnard, 1960, in: (I) placement of
ventral setae on coxa 1 in middle of margin and
not posteriorly; (2) broader coxa 4; (3) distally
tapering article 2 of pereopod 3; (4) much longer
article 2 of pereopod 5; (5) apparent absence of
spine-setae on the outer ramus of uropod 2; (6)
sparseness of setae on epimeron 2.
Phoxocephalus tunggeus appears most closely
similar to P. kergueleni (Stebbing, 1888) but differs
from that species in: (1) presence of posterior setae
on epimeron 3 (but perhaps overlooked on P.
kergueleni); (2) absence of an extra tooth on the
right lacinia mobilis; (3) broader coxa 4; (4) pres-
ence of setae on peduncle of uropod 2; (5) narrower
article 2 of pereopod 4; (6) broader inner plate of
maxilla 2; (7) more distally expanded coxa 1. Sev-
eral important unknown factors of P. kergueleni
include the dactylar apices of pereopods 1-2, the
facial spine formula on article 2 of antenna 2, and
the condition of the inner ramus and the lateral
spination on the outer ramus of uropod 3.
J. L. Barnard (1960:300) presented a table con-
taining a typographical error: "Article 2" of line 3
should read "Article 5." Also, Phoxocephalus
kergueleni is noted in that table as having no thick
spine on article 5 of pereopods 1-2. In retrospect
one may see that this statement was overemphasized
or could be misinterpreted; P. kergueleni has the
spine, it is simply thin, perhaps like that shown
here for P. tunggeus. Our examination of Austral-
ian species of Phoxocephalus suggests that far more
valuable characters will eventually be found to dis-
tinguish the species than those in the table pre-
sented by J. L. Barnard (1960).
MATERIAL.?AM, one sample (1).
DISTRIBUTION.?Victoria, off Cape Everard (now
called Point Hicks), 300 m.
Phoxocephalus rupullus, new species
FICURES 220-222
DESCRIPTION OF FEMALE.?Head about 18 percent
of total body length, greatest width about 67 per-
cent of length, rostrum exceeding apex of article 2
on antenna 1, unconstricted, weakly tapering,
apically rounded. Eyes small-medium in size, clear
in preservative. Article 1 on peduncle of antenna 1
about 1.6 times as long as wide, almost twice as
wide as article 2, apicoventral margin with several
setules, strongly produced dorsal apex with 5
432
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 220.?Phoxocephalus rupullus, new species, holotype, female "a," 7.21 mm.
NUMBER 245 433
El
Dl
FIGURE 221.?Phoxocephalus rupullus, new species, holotype, female "a," 7.21 mm.
434 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
setules; article 2 about half as long as article 1,
apicoventral surface with 4 setae; article 3 attached
weakly to lateral face of article 2 with override of
article 2 medially; accessory flagellum much shorter
than primary flagellum, 3-articulate; primary
flagellum 5-articulate, aesthetascs present. Spine
formula on article 4 of antenna 2 = 1-2-4-2, proxi-
mal groups with accessory setule, distal and disto-
ventral margins with 4 spines widely spread, dorsal
margin with group composed of setule, spine and
long seta, ventral margin with 4 subbasal penicillate
setules, 2 faciolateral setules, and 3 groups of 2-3
setae each; article 5 about 0.6 times as long as
article 4, facial spine formula = 2-2, with one
proximal accessory setule, dorsal margin smooth,
ventral margin with 2 sets of 1-2 setae each, disto-
ventral corner with 2 long spine-setae and setule;
flagellum almost as long as article 4 of peduncle,
5-articulate. Palpar hump of mandible medium;
right incisor weakly tridentate, weakly webbed be-
tween two proximal teeth, left basically bidentate;
right lacinia mobilis flabellate, distal branch poorly
defined, margin toothed; left lacinia mobilis flabel-
late, 5-toothed; right rakers 2 and some rudimen-
taries, left 3 and some rudimentaries; molar tritura-
tive, on medium length pedicle; palp article 2 with
pair of small basal setae and 5 long inner marginal
setae in tandem, article 3 about as long as article 2,
oblique apex with 10 setae. Lower lip with no ap-
parent cones on outer lobe but with 4-5 emergent
salivary ducts, mandibular lobes of medium length,
weakly attenuate and apically rounded, inner lobes
abutted and appearing separate for much of their
length, perhaps raphe extremely distinct. Inner
plate of maxilla 1 large, elliptical, naked; outer
plate with 7 spines; palp failing to reach apex of
outer plate, bearing 2 apical, one apicolateral, one
apicomedial setae almost strap-shaped. Inner plate
of maxilla 2 broader than but as long as outer
plate, medial margin of inner plate with 4 small
setae, none apical, outer plate with 6 apical and 4
medial setae. Inner plate of maxilliped with one
mediofacial seta, 0-1 medial seta, 2-3 apical spines
and one apical setule; outer plate with 3 spines;
article 2 of palp weakly setose medially, article 3
with 3 facial setae, article 4 short, apical spine long
relative to article 4, bearing 2 accessory setules.
Coxa 1 distally expanded, rounded but subacumin-
ate anteroventrally, long setal formula of coxae
1-4 = 9-8-8-6, posteriormost seta of coxae 1-3
short; anterior and posterior margins of coxa 4
divergent, posterior margin almost straight, postero-
dorsal corner subsharp, posterodorsal excavation of
medium size and L-shaped, ratio of width to length
= 1:1. Gnathopod 2 much stouter than gnathopod
1, long posterior setae on article 2 of gnathopods
1-2 = 3 and 3, on both gnathopods article 5 small
and cryptic, article 6 attached to produced apex of
article 5 but held tightly flexed, article 6 stout,
ovatorectangular, palm oblique, defining hump
large on gnathopod 2, apices of dactyls with flake,
length ratios of articles 5-6 on gnathopods 1-2 =
28:72 and 32:110, width ratios = 23:44 and 25:65.
Pereopods 1-2 similar to each other, long pos-
terior setae on article 2 = 3 and 5, article 4
with 5 facial setae, article 5 with 1 and 2 facial
setae, posterior margins of articles 4-5 moderately
setose, article 5 with long distal spine reaching al-
most fully along article 6, with very small con-
tiguous spine at base, article 6 with rows of 3 and 3
or 3 and 2 (pereopod 2) spines, two of these spines
on midposterior margin of article 6, also l)earing
one middistal spinule, spines blunt, dactyl lacking
apical flake, bearing thin inner setule appressed in
marginal groove guarded by protrusion, apical nail
almost fully fused to dactyl. Coxae 5-7 postero-
ventral setule formula = 1-1-3. Width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 52:30:25:14, of
pereopod 4 = 68:29:21:12, of pereopod 5 = 86:
26:17:9, length ratios of articles 2, 4, 5, 6 of
pereopod 3 = 61:27:27:26, of pereopod 4 = 89:
52:46:45, of pereopod 5 = 102:24:28:23; article
2 of pereopod 5 slightly exceeding middle of
article 4; apex of article 6 finely combed. Epimera
1-2 with facial ridge, epimeron 1 with rounded-
quadrate posteroventral corner, epimeron 2 broadly
rounded posteroventrally, posterior margins con-
vex, epimeron 1 with one ventral seta, epimeron 2
with crescentic row of 8 facial setae and one super-
numerary facial seta, epimeron 3 with rounded pos-
teroventral corner, posterior margin convex, bear-
ing setal notches and 6 posterior setae, postero-
ventral margin with 2 setae in notches, epimera 1-2
with small dorsoposterior setule. Urosomite 1 bear-
ing 2 setules on lateral face at base of uropod 1,
urosomite 2 with one such seta, one dorsolateral
seta on each side of urosomite 1, urosomite 3 with
long seta at ventral base of uropod 3. Peduncle of
uropod 1 lacking basofacial setae (see urosomite 1),
apicolateral corner of peduncle with one spine,
NUMBER 245 435
FIGURE 222.?Phoxocephalus rupullus, new species, holotype, female "a," 7.21 mm.
medial margin with 3 spines; rami of uropods 1-2
with apical nail almost fully amalgamated with
ramus, outer rami of uropods 1-2 and inner ramus
of urojxxl 1 with one dorsal spine each, inner ramus
of uropod 1 with one mediobasal setule, outer
ramus of uropod 1 as long as or longer than inner
ramus, almost straight, rami and peduncles of uro-
pods 1-2 lacking combs. Uropod 3 long, rami
feminine, peduncle with 3 ventrolateral short
spines, medially with setule, dorsolaterally with
spinule, inner ramus extending to M. 43 on article
1 of outer ramus, apex with one seta, article 2 of
outer ramus short, 0.17, bearing 2 long apical setae,
outer margin of article 1 with 4 acclivities, setal
formula = 1-1-2-2-2, one seta of some pairs short,
medial apex with one long seta. Telson ordinary,
length-width ratio = 1:1, apices scarcely tapering,
broadly rounded, each with weak lateral acclivity
bearing one long seta, one short lateral seta, these
separated by shorter plusetule, dorsal pairs of
plusetules diverse. Cuticle covered with knobs and
villi, these knobs on posterior coxae and article 2
of pereopods 3-5 organized into polygons.
ILLUSTRATIONS.?Upper lip drawn from slightly
oblique dorsal view; inner lobes of lower lip seen
imperfectly at base; dactyl of pereopod 4 on both
sides missing.
HOLOTYPE.?AM, female, "a," 7.21 mm. Unique.
[Extremely large, possibly senile but brood plates
bearing setae, thus indicating presenility.]
TYPE-LOCALITY.?AM P. 18248, 18 Jun 1962, off
Syndey, New South Wales, Australia, 3S?58.4'S,
151 ^ E , 150 m.
RELATIONSHIP.?This species bears similarities to
Phoxocephalus regium K. H. Barnard, 1930 (see
Hurley, 1954), from New Zealand. In P. regium
only 3 setae occur on epimeron 3 and the horizontal
row of setae on epimeron 2 is confined to the an-
terior part of the plate; P. regium lacks a dorsal
spine on the outer ramus of uropod 1 and no spines
on the rami of uropod 2. The inner ramus of uro-
pod 3 is elongate. Article 4 of pereopod 5 is thin in
P. regium, articles 4-5 of pereopods 3-4 are even
thinner than those of P. rupullus, gnathopod 1 ap-
pears much larger than in P. rupullus and article 5
longer on P. regium than in P. rupullus. Coxa 4 has
a sharper posterior lobe in P. rupullus, longer
apical nail on the dactyl of the maxilliped, and
lacks a basofacial seta on article 3 of the mandibu-
lar palp.
MATERIAL.?AM, one sample (1).
DISTRIBUTION.?New South Wales, off Sydney,
150 m.
Phoxocephalus burletts, new species
FICURES 223-225
DESCRIPTION OF FEMALE.?Head about 18 percent
of total body length, greatest width about 55 per-
cent of length, rostrum nearly reaching middle of
article 2 on antenna 1, unconstricted, weakly taper-
436
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 223.?Phoxocephalus burleus, new species, holotype, female "f," 5.93 mm (n ? male
"n," 4.54 mm).
NUMBER 245 437
ing, apically rounded. Eyes of medium size, dark
core surrounded by one partial layer of clear
ommatidia in preservative. Article 1 on peduncle of
antenna 1 about 1.8 times as long as wide, 1.8 times
as wide as article 2, apicoventral margin with sev-
eral setules, strongly produced dorsal apex with 6
setules; article 2 about 0.42 times as long as article
1, apicoventral surface with 4 setae; article 3 essen-
tially attached terminally to article 2; accessory
flagellum shorter than primary flagellum, S-articu-
late; primary flagellum 5-articulate. aesthetascs ab-
sent. Spine formula on article 4 of antenna 2 =
3-4-3, proximal sets with 1-2 accessory setules,
spines thin, dorsal margin with one short spine, one
long seta, ventral margin with 3 subbasal penicillatc
setules and 10 long or short setae, apex with 4
spines; article 5 less than 0.6 times as long as article
4, facial spine formula = 2-2. with one proximal
accessory setule, dorsodistal corner with one spine-
seta, apex with one spine, ventral margin with 2
pairs of long-short setae, ventrodistal apex with one
long, one short setae; flagellum about as long as
article 4 of peduncle. 5-articulate. Palpar hump of
mandible small; right incisor weakly tridentate,
with weak accessory tooth between teeth 2 and 3;
left basically bidentate, distal branch weakly tri-
dentate; right lacinia mobilis flabellate, distal
branch poorly defined, margin toothed; left lacinia
mobilis flabellate, irregularly toothed; right rakers
2, left 3; molar triturative, on long pedicle; palp
article 2 with 5 long medial setae on distal half,
article 3 about as long as article 2, oblique apex
with 10 setae. Mandibular lobes of lower lip of
medium length, weakly attenuate and apically
rounded, inner lobes fused together for most of their
length but with distinct raphe. Inner plate of
maxilla 1 large; elliptical, naked, outer plate with
7 spines; palp scarcely exceeding apex of outer
plate, bearing 2 apical and one apicolateral setae.
Inner plate of maxilla 2 scarcely broader than and
extending equally with outer plate, medial margin
of inner plate with 2 setae, apical margin with 3
setae; outer plate with 2 apicolateral, 3 apical and
3 medial setae. Inner plate of maxilliped with one
mediofacial seta, 2 apical spines; outer plate with
4-5 spines; article 2 of palp with sparse medial
setae, article 3 with 2-3 facial setae, article 4 of
medium length, apical spine long, bearing 2 acces-
sory setules. Coxa 1 distally expanded and narrowly
rounded anteroventrally; long setal formula of
coxae 1-4 = (8-10)-(9-I0)-(8-10)-(5-8), posterior-
most seta of coxae 1-3 shortened (= elongate
setule); anterior and posterior margins of coxa 4
divergent, posterior margin weakly convex, postero-
dorsal corner subsharp, posterodorsal excavation of
medium size and V-shaped, ratio of width to length
= 73:86. Gnathopod 2 much stouter than gnatho-
pod 1; long posterior setae on article 2 of gnatho-
pods 1-2 = 2 and 5, short anterior setae = 1 and 3;
article 3 short, only gnathopod 2 weakly eusirid,
article 6 broadened but elongate, rectangular, palm
oblique, defining hump large on both pairs, dactyl
fitting palm, apices of dactyls with flake, posterior
margin of article 5 on gnathopod 1 free, on gnatho-
pod 2 cryptic; length ratios of articles 5-6 on
gnathopods 1-2 = 34:77 and 35:126, width ratios
= 22:43 and 18:66. Pereopod 2 slightly thinner
than pereopod 1; long posterior setae on article 2
of pereopods 1-2 = 2-3 and 3, short anterior setae
= 2 and 1; article 4 of pereopods 1-2 with 6 and 2
subapical anterior setae, posterior margins of article
4 moderately setose, of article 5 more densely so,
article 5 with 3 and 2 anterofacial setae, with long
posterodistal spine reaching to M. 95 on article 6,
with small contiguous spine at base, article 6 with
one long posterior setal-spine and rows of 3 and 3
spines and setae, plus middistal spine, some stout,
some thin, dactyl lacking apical flake, inner margin
with acclivity and tooth bearing setule appressed in
wide groove, apical nail fused almost fully to dactyl.
Coxae 5-7 posteroventral setule formula = 1-1-1.
Width ratios of articles 2, 4, 5, 6 of pereopod 3 =
55:25:21:12, of pereopod 4 = 72:26:18:10, of pereo-
pod 5 = 89:21:17:8, length ratios of pereopod
3 = 76:24:26:26, of pereopod 4 = 98:57:56:47, of
pereopod 5 = 112:20:28:22; article 2 of pereopod
5 slightly exceeding apex of article 4, posterior
margin with accessory tightly packed group of
dorsoposterior setules; apex of article 6 finely
combed. Epimera 1-2 with facial ridge, each
with rounded-quadrate posteroventral corner and
convex posterior margin, epimeron 3 with rounded
posteroventral margin, posterior margin broadly
convex, bearing long setae set in notches, margins
between notches inflated, setae extending onto
postroventral margin, epimeron 1 with one postero-
ventral seta, epimeron 2 with 5-7 facial setae in
horizontal row. Urosomite 1 bearing 2 short lateral
setae at base of uropod 1; urosomite 2 with one
such seta; urosomites dorsally truncate; urosomite
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 224.?Phoxocephalus burleus, new species, holotype, female "f," 5.93 mm (n ?? male
"n," 4.54 mm).
NUMBER 245 4S9
3 with 2 long setae at ventral base of uropod 3.
Peduncle of uropod 1 with one setule not strongly
basal, lateral apex of peduncle with one spine,
medial margin with 4 spine-setae; rami of uropods
1-2 with apical nails scarcely articulate but con-
spicuous, outer ramus of uropods 1-2 with 2 dorsal
spines (elongate), inner ramus of uropod 1 with one
spine, of uropod 2 naked, inner ramus of both
uropods 1-2 with mediobasal spinule (unusual for
uropod 2), outer ramus of uropod 1 slightly longer
than inner, of normal dimensions and shape; lateral
apex of peduncle on uropod 2 with stout spine,
margin with 4 long setae, apicomedial corner of
peduncle with long spine-seta: comb on apices of
peduncles of uropods 1-2 obsolescent. Uropod 3
long, rami feminine, peduncle with 4 long ventro-
lateral spine-setae, dorsolaterally with tiny spinule,
dorsomedially with tiny spinule, inner ramus ex-
tending to M. 60 on article 1 of outer ramus, apex
with one seta, article 2 of outer ramus elongate,
0.37, bearing 2 long apical setae, outer margin of
article 1 with 3 acclivities, elongate spine-seta for-
mula ? 1-1-1-3, medial apex with one spine-seta.
Telson slightly elongate, length-width ratio about
14:11, apices scarcely tapering, each with medial
protrusion and lateral acclivity bearing one long
spine-seta and one short lateral setule, dorsal pair
of plusetules diverse. Cuticle armed sparsely with
bulbar setules and densely covered with fingerprint
striations heavily cornified and thickened.
DESCRIPTION OF MALE (male "n," 4.54 mm,
unique specimen).?Eyes enlarged but only of
medium size. Article 1 of antenna 1 with medial
patch of fuzz; articles 1-2 of primary flagellum
each with calceolus and one large aesthetasc. An-
tenna 2 enlarged, elongate; dorsomedial margins of
articles 3 and 4 with fuzz; facial spine formula of
article 4 = 3-3-3 plus 3 other apical spines, only
one accessory setule with each proximal spine
group, setae of articles 4-5 fewer and shorter than
in female, article 5 with 3 dorsal sets of male setae
and one calceolus; flagellum elongate, flagellar
formula = (20), 1-2 3, 5, 7 . . . 19. Long posterior
setae on article 2 of gnathopods 1-2 and pereopods
1-2 = 1-2-2-3, short anterior setae = 1-1-1-1. Arti-
cle 2 of pereopods 4?5 narrower and more elongate
than in female. Epimera 1-3 broadened anteropos-
teriorly, (epimeron 1 poorly preserved but of di-
mensions commensurate with epimeron 2, presence
of one seta undetermined); epimeron 2 with 5 facial
setae; setae on epimeron 3 shorter than in female.
Setules on urosomites 1-2 at bases of uropods elong-
ate, urosomite 1 bearing only one setule. Spines
and setae on uropods 1-2 shorter than in female,
peduncle of uropod 2 with 9 short lateral spines
not divisible into size classes, medial margin with 3
spines, peduncle of uropod 1 with 2 apicolateral
spines, otherwise spine formula of uropods 1-2 same
as in female. Uropod 3 masculine, inner ramus
extending to apex of outer ramus on article 2, both
rami setose on both margins, lateral margin on
article 1 of outer ramus with 4 acclivities, formula
of long setae = 1-1-1-1-1, formula of short setae
= 1-1-1-1-2; article 2 of outer ramus shorter than
in female, spines on peduncle shortened, apico-
medial margin of peduncle with 3 setules in 2
groups. Telson of dimensions similar to female,
apical spines short, each lobe dorsomedially with
row of denticles.
VARIATIONS.?Female "h," 4.53 mm, much
smaller than other females analyzed, with 7 setae
on epimeron 2 and 5 setae plus one spine on lateral
margin of peduncle on uropod 1, highest counts
found in females of the species.
HOLOTYPE.?AM, female "f," 5.93 mm.
TYPE-LOCALITY.?SBS CXS4, 22 May 1972, off
Malabar, New South Wales, Australia, 92 m, sandy
gravel.
VOUCHER MATERIAL.?Type-locality, female "g,"
5.31 mm; SBS D3S3, male "n," 4.54 mm (illus.); SBS
C7S4, female "h," 4.53 mm.
RELATIONSHIP.?This species has its closest af-
finities with P. rupullus, new species. The latter is
based on a giant, perhaps senile female. Some of
the character differences between the two species
may be based on the senilic condition of P. rupullus
but the extraordinary difference between the cuticles
of the two species confirms their distinction.
Phoxocephalus rupullus has villi organized into
polygons whereas P. burleus has thickened striations
lacking a pattern. Phoxocephalus rupullus has more
setae and spines in certain places and fewer in
other places than does P. burleus; because senile
individuals may both gain or lose armaments selec-
tively we leave the reader to compare our illustra-
tions for these differences.
Phoxocephalus regium K. H. Barnard, 1930 (see
Hurley, 1954), from New Zealand, is in the species
group of P. burleus because of equally extending
rami on uropod 1 but otherwise is remote morpho-
440
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 225.?Phoxocephalus burleus, new species, holotype, female "(," 5.93 mm (n ? male
"n," 4.54 mm).
NUMBER 245 441
logically; among numerous distinctions of P. regium
from P. burleus and P. rupullus we emphasize the
following: the confinement of facial setae on epi-
meron 2 to the far anterior region, the shortness of
posterior setae on epimeron 3, the absence of pos-
terior setae on article 2 of pereopod 4, and the ab-
sence of a dorsal spine on the inner ramus of
uropod I.
MATERIAL.?SBS, 3 samples (4).
DISTRIBUTION.?New South Wales, off Malabar,
71-92 m, sandy gravel, medium fine sand.
Phoxocephalus keppeli, new species
Phoxocephalus bassi.?K. H. Barnard, 1930:330-331, fig. 9b
(not Subbing, 1888).
DIAGNOSIS.?Phoxocephalus from New Zealand
with closely packed submarginal bundle of 3 setae
on posterior face of epimeron 3, true posterior edge
of epimeron 3 broadly rounded and naked. Named
in honor of Dr. Keppel H. Barnard.
HOLOTYPE.?Male, 7.5 mm, fig. 9b of K. H.
Barnard, 1930, specimen apparently deposited in
British Museum (Natural History).
TYPE-LOCAI.ITY.?"Discovery" Sta. 110, off Three
Kings Islands, New Zealand, 6 Aug 1911, surface,
night.
REMARKS.?K. H. Barnard described nothing of
this species except epimeron 3 plus comments inti-
mating its resemblance to P. bassi Stebbing (1888).
Barnard suggested that Stebbing overlooked the
bundle of 3 setae on epimeron 3 but we, having
found a specimen closely matching that of Steb-
bing's species near his type-locality, doubt that sup-
position. J. L. Barnard (1972b) found few benthic
amphipods of New Zealand conspecific with those
from Australia.
DISTRIBUTION.?Northern New Zealand, surface.
Phoxocephalus species
NOTES.?A very distinctive and unusual species
but damaged and not worthy of specific description;
true generic placement unknown. Mandibles undis-
sected but molar seen to be solid, conical, possibly
minutely triturative; maxilla 1 with uniarticulate
palp; article 2 of antenna 1 with strong conical
medial protrusion overriding and exceeding apex
of article 3; flagellum of antenna 2 with 3 articles,
article 5 of peduncle elongate (thus like Joubi-
nella); gnathopods stout, article 6 of gnathopod 1
longer than wide, anterior and posterior margins
weakly divergent, palm subtransverse, article 5 mod-
erately elongate; gnathopod 2 slightly larger than
gnathopod 1, article 6 scarcely longer than wide;
article 6 of pereopods 1-2 with spine rows of 1 + 1
plus middistal spine, longest spine shorter than
dactyl; outer ramus of uropod 1 slightly shortened
and curved, apices of rami on uropods 1-2 bluntly
conical apically, with small scales, rami lacking
dorsal spines; uropod 3 short, stubby, outer ramus
with elongate article 2 bearing 2 long setae (atypical
of Phoxocephalus), inner ramus naked, about three-
fourths as long as article 1 of outer ramus; each
apex of telson with one spine towards medial side,
apicolateral setule shorter than spine, dorsal pairs
of plusetules near M. 75 and extremely elongate (a
probable mark of youth).
MATERIAL.?SBS Long Reef, juvenile, 1.95 mm.
DISTRIBUTION.?New South Wales, Long Reef, N
of North Head, Sydney, 43 m.
Phoxocephalus holbolli (Kr0yer)
Phoxus Holbolli Kr0yer, 1842:151-152; 1845:551-563.?Bru-
zelius, 1859:68.?Smith, 1883a:221; 1883b:228.
Phoxus Holbolli.?Chcvreux, 1887:298.
Phoxocephalus holbolli.?]. L. Barnard, 1960:294-298, pis. 47.
48 [with references].
Phoxocephalus holbolli.?Hamond, 1967:118-119.
Phoxocephalus holbolli.?Bousfield, 1973:123-124, pi. 25.1.
DISTRIBUTION.?North and eastern Atlantic from
Greenland south to Plymouth, England; into the
Baltic Sea to east of Bornholm, Barents Sea, White
Sea, Kara Sea; western Atlantic from Greenland
south to Long Island Sound; 0-190 m.
Phoxocephalus homitis J. L. Barnard
Phoxocephalus homilis J. L. Barnard, 1960:301-303, pis. 49.
50; 1964c:245; 1966a:90; 1966b:30.
DISTRIBUTION.?Monterey Bay, California to
Bahia de San Cristobal, Baja California, 60-2059 m.
Phoxocephalus kergueleni (Stebbing)
Phoxus kergueleni Stebbing, 1888:816-819, pi. 55.
Phoxocephalus kergueleni.?Stebbing, 1906:135.?J. L. Barn-
ard, 1960:300; 1964b:21-22, fig. 17; 1967:138, fig. 67.
442 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DISTRIBUTION.?Off Cumberland Bay, Kerguelen,
220 m; eastern Pacific Ocean from Panama Basin
north to Cedros Trench, Baja California, 842-2398
m.
Phoxocephalus reghtm K. H. Barnard
Phoxocephalus regium K. H. Barnard, 1930:351-332, fig. 9a.
?Hurley, 1954:594-598, figs. 68-91.?J. L. Barnard, 1960:
299-300.
Phoxocephalus bassi.?Thomson, 1902:463 [not Stebbing,
1888].
Phoxocephalus kergueleni.?Chilton, 1909:618-619 [not
Stebbing, 1888].
DISTRIBUTION.?New Zealand, off Three Kings
Islands, Otago Harbour; Snares Islands; surface to
91 m.
Phoxocephalus tenuipes Stephensen
Phoxocephalus tenuipes Stephensen, 1925:160-161, fig. 47.?
J. L. Barnard, 1960:298.
DISTRIBUTION.?West of Greenland, 66?35'N;
North Atlantic, west and southwest of Iceland;
600-1505 m.
Jerildaria, new genus
DIAGNOSIS.?Eyes present. Flagellum of antenna 2
reduced in female. Article 2 of antenna 1 ordinary,
ventral setae confined apically. Article 1 of antenna
2 not ensiform; article 3 with 2 setules; facial spines
on article 4 in 2 or more rows, plus special apical
spines; article 5 ordinary in size. Right mandibular
incisor with 3 teeth; molar strongly triturative,
bearing fuzz; palpar hump medium to large. Palp
of maxilla 1 uniarticulate, inner plate naked. Seta-
tion of maxilla 2 ordinary. Inner plate of maxilli-
peds ordinary, apex of palp article 3 not protuber-
ant, dactyl elongate, apical nail distinct, medium.
Gnathopods dissimilar, weakly to moderately en-
larged; article 5 of gnathopod 2 of ordinary length,
almost cryptic, elongate on gnathopod 1, with
eusirid attachment, palms oblique to transverse,
hands of gnathopods 1-2 rectangular, broadened,
poorly setose anteriorly. Article 5 of pereopods 1-2
setose posteroproximally. Article 2 of pereopod 3 of
broad form, articles 4-5 of pereopods 3-4 narrow
to medium, article 2 of pereopods 3-4 not setose
posteriorly; pereopod 5 ordinary, article 2 naked
ventrally, article 3 ordinary, dactyl normal. Pedun-
cle of uropod 1 normally elongate, without apico-
ventral spike, without special enlarged apicolateral-
medial spine, peduncular apices of uropods 1-2
combed, inner ramtis of uropod 1 with marginal
spines in one row, no rami continuously spinose to
apex, inner ram us of uropod 2 ordinary. Uropod
3 ordinary, short article 2 of outer ramus carrying
2 long apical setae. Telson elongate, with 2-4
apical spines or setae on each lobe, without special
dorsal and lateral spines or setae. Kpimera 1-2
bearing few long posterior setae, without midfacial
setae above ventral facial ridge; epimeron 3 ordi-
nary. Urosomite 1 generally naked except for
sparse apicoventral setae or spines near base of
uropod 1, urosomite 3 without dorsal hook or
special process.
DESCRIPTION?Rostrum fully developed. [Fuzz on
article 1 of antenna 1 in male unknown, calceoli
on male antennae 1-2 unknown.] Prebuccal parts
ordinary, weakly extended forward, distinct, upper
lip dominant. Right lacinia mobilis bifid, flabel-
late; mandibular palp medium to thin, article 1
short, article 2 without outer setae, apex of article
3 oblique. Lower lip bearing cones. Outer plate
of maxilla 1 with 9 spines, no spine especially
thickened. Inner plates of maxilliped thick, ordi-
narily setose. Coxae 2-4 without special antero-
dorsal humps. All posterior spines on article 6 of
pereopods 1-2 thick and stiff, midapical spine or
seta present. Article 2 of pereopod 5 without facial
setae. Peduncle of uropod 1 with dorsolateral spines
confined apically, medial spines confined apically;
peduncle of uropod 2 with only one medial spine
confined apically. Peduncle of uropod 3 lacking
extra subapical setae or spines. Telson with ordi-
nary pair of midlateral or dorsal setules on each
side highly medial.
TYPE-SPECIES.?Jerildaria joubiphoxus, new spe-
cies.
COMPOSITION?Unique.
RELATIONSHIP.?The superficial and external ap-
pearance of this genus suggests relationship to the
pelagic Joubinella but Jerildaria has only one arti-
cle in the first maxillary palp, has gnathopod 2
larger than gnathopod 1, lacks dorsodistal spines
and freely articulate nails on the rami of uropods
1-2, and the palms of the gnathopods do not pro-
trude as in Joubinella. Distinctive generic char-
NUMBER 245 443
acters of Joubinella also include an elongation of
article 4 on pereopod 4, a strongly shortened arti-
cle 4 of pereopod 5, and the absence of strong lat-
eral spination on article 4 of antenna 2.
Jerildaria differs from Phoxocephalus, also bear-
ing triturative molar, in only one strong character,
the greatly enlarged gnathopods with elongate,
eusirid-like article 5 of gnathopod 1. The extended
palpar base on the mandibular body of Jerildaria
is uncharacteristic of Phoxocephalus as are the
following characters: the short article 2 on the
outer ramus of uropod 3, the thick rami of uropods
1-2, the slightly elongate article 5 of antenna 2,
and the extreme shortness of the inner ramus on
uropod S.
Jerildaria differs from the primitive Ponthar-
pinia in the loss of articulation on the first maxil-
lary palp and in the absence of dorsodistal spines
on the rami of uropods 1-2. The latter character
is not so vital because species in Phoxocephalus
have these spines present or absent. In Pontharpinia
the body of the mandible is more strongly ex-
tended to the base of the palp than in Jerildaria
and the gnathopods are much smaller and not
eusirid, though gnathopod 2 of Pontharpinia is
enlarged.
Jerildaria joubiphoxus, new species
FIGURES 226, 227
DESCRIPTION (female-like).?Head about 18 per-
cent of total body length, greatest width about 64
percent of length; rostrum unconstricted, broad,
short, almost reaching apex of article 2 on antenna
1. Eyes medium, largely occluded with pigment.
Article 1 on peduncle of antenna 1 about 1.2 times
as long as wide, about 2.1 times as wide as article
2, apicoventral margin with about 8 setules,
strongly produced dorsal apex with 4 setules; arti-
cle 2 about 0.85 times as long as article 1, with
apicoventral row of 3^t setae; primary flagellum
widi 5 articles, about 0.75 times as long as pe-
duncle, bearing several short aesthetascs; accessory
flagellum elongate, with 4 articles. Spine formula
on article 4 of antenna 2 = 2-4-4 or 2-4-5, one
spine of proximal group thin, all others thick,
dorsal margin with notch bearing one seta and one
spine, ventral margin with 3 groups of 1-2 medium
setae, 2 ventrodistal short thick spines and 2 mid-
distal long spines; article 5 about 0.8 times as long
as article 4, facial spine formula = 2-2, dorsal mar-
gin naked, ventral margin with one set of setae,
ventrodistal medium seta, setule and spine; flagellum
about as long as article 5 of peduncle, with 4-5
articles. Mandibles with medium palpar hump;
right incisor with 3 teeth; left incisor with 2 weak
branches; right lacinia mobilis bifid, distal branch
little longer than proximal, flabellate, broad, den-
ticulate, with one facial hump, proximal branch
simple, pointed; left lacinia mobilis with 4 teeth
plus weak accessory teeth, middle teeth shortened;
right rakers 2; left rakers 3; molars triturative; palp
article 1 short, article 2 with 2-3 long inner apical
setae and one other shorter inner basal seta, article
3 about 0.85 times as long as article 2, oblique apex
with 8 spine-seta, basofacial formula = 0. Inner
plate of maxilla 1 ordinary, naked, palp with 2
apical setae, one apicomedial and one apicolateral
setae. Plates of maxilla 2 extending subequally,
equally wide, outer with one apicolateral seta,
inner with 11 medial setae. Inner plate of maxilli-
ped with 3 thin apical spines and setae, 2-3 medio-
facial setae, outer plate with 5-6 medial and apical
spines, basalmost thin, medial margin of article 2
moderately setose, article 3 not protuberant, with
5 facial setae, no lateral setae, nail of article 4
medium, with 2 accessory setules. Coxa 1 expanded
distally, anterior margin straight; main ventral
setae of coxae 1 ^ = (4-5)-(4-5)-(4y-(l-2), poste-
riormost seta of coxae 1-3 shortest; anterior and
posterior margins of coxa 4 strongly divergent, pos-
terior margin very oblique, almost straight, postero-
dorsal corner subsharp, posterodorsal margin L-
shaped, width-length ratio of coxa 4 = 1:1. Long
posterior setae on article 2 of gnadiopods 1-2 and
pereopods 1-2 = 0-3-(2-3)-3, short anteriors =
2-2-{2-3)-2, no others; gnathopods with stout sixth
article, that of gnathopod 2 larger; width ratios of
articles 5-6 on gnathopods 1-2 = 20:70 and 21:73,
length ratios = 88:62 and 64:80; palmar humps
small, palms almost transverse; article 5 of gnatho-
pod 1 elongate, elliptical, thin, posterior margin
weakly rounded, long; article 5 of gnathopod 2
elongate, elliptical, thin, posterior margin rounded,
short. Pereopods 1-2 similar; posterior margin of
article 4 on pereopod 2 slightly more setose than
on pereopod 1, facial setae formula on article 4 =
1-1, on article 5 = 0-0; main spine of article 5
extending to M. 70 on article 6, article 5 with no
444 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 226.?Jerildaria joubiphoxus, new species, holotype, sex unknown "b," 5.50 mm.
NUMBER 245
FIGURE 227.?Jerildaria joubiphoxus, new species, holotype, sex unknown "b," 5.50 mm (w
sex unknown "w," 5.46 mm).
446 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
proximoposterior spines; spine formula of article
6 = 2 + 2 plus middistal spine plus posterior spine,
some spines short; acclivity on inner margin on
dactyls of pereopods 1-2 weak, emergent setule
long, midfacial pluseta very large, highly proximal,
nail weakly distinct. Coxae 5-7 posteroventral set-
ule formula = 1-1-2. Articles 4-5 of pereopods 3-4
narrow, facial spine rows sparse, pereopod 3 espe-
cially small, facial ridge formula on article 2 of
pereopods 3-5 = 0-1-1; width ratios of articles 2,
4, 5, 6 of pereopod 3 = 44:21:19:11, of pereopod
4 = 66:27:21:12, of pereopod 5 = 90:20:18:10,
length ratios of pereopod 3 = 52:19:29:30, of
pereopod 4 = 82:50:45:34, of pereopod 5 = 1 1 0 :
24:30:28; article 2 of pereopod 5 not reaching
middle of article 5; medial apex of article 6
very finely combed. Posteroventral corner of epi-
meron 1 rounded, posterior margin weakly convex,
margin above corner with setule, anteroventral
margin naked, posteroventral face with one or no
long seta; posteroventral corner of epimeron 2
rounded-quadrate, posterior margin straight, bear-
ing one setule, facial setae = 4, posteriormost pair
set vertically; posteroventral corner of epimeron 3
weakly protuberant, with setule sinus, posterior
margin straight, with 2 long setae set in notches,
occasional additional setule notch, ventral margin
and face naked; epimera 1-2 with large seta on
posterodorsal margin set in weak notch. Urosomite
1 with lateral setule at base of uropod 1, long ven-
tral seta, articulation line complete; urosomite 3
weakly protuberant dorsally. Rami of uropods 1-2
with fused enlarged apical nails, outer ramus of
uropod 1 with 2 dorsal spines, inner with 2, outer
ramus of uropod 2 with one dorsal spine, inner
naked; peduncle of uropod 1 with one small apico-
lateral spine, medially with 3 marginal spines, api-
calmost ordinary; peduncle of uropod 2 with 4
dorsal spines, medially with one small apical spine;
apicolateral corners on peduncles of uropods 1-2
lacking comb. Peduncle of uropod 3 with 2 ventral
spines, dorsally with one lateral setule, one medial
spine; rami feminine, inner extending to M. 33 on
article 1 of outer ramus, apex with one short seta,
article 2 of outer ramus short, 0.10, bearing 2 long
setae, apicomedial margin of article 1 with sets of
2 and 3 setae, lateral margin with 4 acclivities, setal
formula = 1-1-2-3-3. Telson ordinary, length-
width ratio = 35:31, almost fully cleft, each apex
wide, truncate, lateral acclivity weak, bearing lat-
eral setule, spine next medial much longer, medi-
ally with short seta, occasionally with extra lateral
short seta, midlateral setules ordinary to large.
Cuticle with ordinary sparse bulbar setules of vary-
ing sizes mixed with pipes, emergent setules occa-
sionally branched.
OBSERVATIONS.?Epistome protruding in front of
upper lip, both parts fully articulate, upper lip
with weak midhump ventrally, bearing minute
sharp cusps; inner lobes of lower lip fused for mo*l
of length, marked medially with longtitudinal
plaque, each outer lobe with cone. Neither speci-
men of this species shows any mark of sex except
for one possible rudimentary penial process on
specimen "w".
ILLUSTRATIONS.?Outline of epistome shown on
lateral view of head: left eye of holotype displaced
medially in dorsal view of head; left mandible and
article 3 of antenna 1 not flattened; apically
rounded spines on uropods 1-2 blunted by damage.
HOLOTYPE.?AM, sex unknown "b," 5.50 mm.
TYPE-LOCALITY? SBS Long Reef. 27 Apr 1972.
Long Reef, New South Wales, N of North Head,
Sydney, Australia, 43.3 m.
VOUCHER MATERIAL.?Type-Locality, sex un-
known "w," 5.46 mm (illus.), only known specimen
in addition to holotype.
DISTRIBUTION.?New South Wales, Long Reef,
Sydney, 43 m.
Leptophoxoides J. L. Barnard
Leptophoxoides J. L. Barnard, 1962:51.
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 reduced in female. Article 2 of antenna 1 espe-
cially shortened, ventral setae if present confined
apically. Article 1 of antenna 2 not ensifonn;
[? article 3 with 2 setules]; facial spines on article 4
in one main row; article 5 especially short. Right
mandibular incisor with 3 teeth; molar strongly
triturative, not bearing fuzz; palpar hump small.
Palp of maxilla 1 uniarticulate; inner plate naked.
Setation of maxilla 2 weak. Inner plate of maxilli-
peds ordinary; apex of palp article 3 strongly pro-
tuberant, dactyl elongate, apical nail distinct, me-
dium. Gnathopods dissimilar, gnathopod 2 strongly
enlarged; article 5 of gnathopods 1-2 very short,
cryptic, with eusirid attachment, palms oblique,
hands of gnathopods 1-2 ordinary to broadened,
NUMBER 245 447
poorly setose anteriorly. Article 5 of pereopods 1-2
setose posteroproximally. Article 2 of pereopod 3
of broad form, articles 4-5 of pereopods 3-4 nar-
row, article 2 of pereopods 3-4 not setose poste-
riorly, pereopod 5 ordinary, article 2 naked ven-
trally, article 3 ordinary, [? dactyl normal]. Epimera
1-2 lacking long posterior setae, without midfacial
setae above ventral facial ridge, epimeron 3 of
rounded classification and lacking long setae. Uro-
somite 1 generally naked; urosomite 3 without dor-
sal hook or special process. Peduncle of uropod 1
normally elongate, without apicoventral spike, with-
out special enlarged apicolateral-medial spine, only
peduncular apices of uropod 1 combed, inner ramus
of uropod 1 with marginal spines in one row, no
rami continuously spinose to apex, inner ramus of
uropod 2 ordinary. Uropod 3 ordinary; elongate
article 2 of outer ramus carrying one medium apical
seta. Telson elongate, with only one apical spine on
each lobe plus setules, without special dorsal and
lateral spines or setae.
DESCRIPTION.?Rostrum fully developed. [Fuzz
on article 1 of antenna 1 in male unknown, cal-
ceoli on male antennae 1-2 unknown. Prebuccal
parts unknown.] Right lacinia mobilis [? bifid or
simple], thin; article 1 of raandibular palp short,
palp medium to thick, article 2 without outer setae,
apex of article 3 oblique. Lower lip lacking cones.
Outer plate of maxilla 1 with 7 spines, no spine
especially thickened. Inner plates of maxilliped
especially thin, poorly armed. Coxae 2-4 without
special anterodorsal humps. All posterior spines
on article 6 of pereopods 1-2 thin and seta-like,
[midapical spine or seta unknown]. Article 2 of
pereopod 5 without facial setae. Peduncle of uro-
pod 1 with dorsolateral spines widely spread, me-
dial spines confined apically; peduncle of uropod
2 with only one medial spine or setule confined
apically. Peduncle of uropod 3 lacking extra sub-
apical setae or spines. Telson with ordinary pair
of midlateral or dorsal setules on each side.
TYPE-SPECIES.?Leptoph.oxoid.es molaris J. L.
Barnard, 1962 (monotypy).
COMPOSITION.?Unique.
Leptophoxoides molaris J. L. Barnard
Leptophoxoides molaris J. L. Barnard, 1962:51, fig. 40.
DISTRIBUTION.?Cape Basin, off South Africa,
4961m.
Leptophoxus San
Leptophoxus San, 1895:146.
DIAGNOSIS.?Eyes absent. Flagella of antennae
1-2 reduced in female. Article 2 of antenna 1 espe-
cially shortened, ventral setae confined apically.
Article 1 of antenna 2 not ensiform; [Particle 3
with 2 setules]; facial spines on article 4 in 2 rows;
article 5 ordinary in size. Right mandibular incisor
with 3 teeth; molar not triturative, small, pillow-
shaped, bearing semiarticulate spines, [Pusually not
bearing fuzz]; palpar hump medium. Palp of max-
illa 1 uniarticulate; inner plate naked. Setation of
maxilla 2 weak. Inner plate of maxillipeds ordi-
nary, apex of palp article 3 strongly protuberant,
dactyl elongate, apical nail distinct, short. Gnatho-
pods dissimilar, gnathopod 2 strongly enlarged;
article 5 of gnathopods 1-2 very short, cryptic on
gnathopod 2, with eusirid attachment, palms
oblique, hand of gnathopod 2 broadened, both
hands poorly setose anteriorly. Article 5 of pereo-
pods 1-2 setose posteroproximally. Article 2 of
pereopod 3 of broad form, articles 4-5 of pereopods
3-4 narrow, article 2 of pereopods 3-4 not setose
posteriorly; pereopod 5 ordinary, article 2 naked
ventrally, article 3 ordinary, dactyl normal. Epi-
mera 1-2 lacking long posterior setae, without mid-
facial setae above ventral facial ridge; epimeron 3
of rounded classification and lacking long setae.
Urosomite 1 generally naked; urosomite 3 without
dorsal hook or special process. Peduncle of uropod
1 normally elongate, without apicoventral spike,
without special enlarged apicolateral-medial spine;
[Ppeduncular apices of uropods 1-2 not combed];
inner ramus of uropod 1 widi marginal spines in
one row, no rami continuously spinose to apex,
inner ramus of uropod 2 ordinary. Uropod 3 ordi-
nary, elongate article 2 of outer ramus carrying 2
short apical setae. Telson elongate, with only 2
apical spines or setae on each lobe plus setules,
without special dorsal and lateral spines or setae.
DESCRIPTION.?Rostrum fully developed. Fuzz
present on article 1 of primary flagellum on male
antenna 1; [fuzz on article 1 of antenna 1 in male
unknown; calceoli on male primary flagellum of
antenna 1 unknown]. Calceoli on article 5 of male
antenna 2 present; flagellum in male with calceoli.
Prebuccal parts ordinary, poorly separated from
each other, upper lip dominant. Right lacinia mo-
bilis bifid, thin, article 1 of mandibular palp me-
448 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
dium to thick, article 1 short, article 2 without
outer setae, apex of article 3 oblique. Lower lip
lacking cones. Outer plate of maxilla 1 with 7
spines, no spine especially thickened. Inner plates
of maxilliped especially thin, poorly armed. Coxae
2-4 without special anterodorsal humps. All pos-
terior spines on article 6 of pereopods 1-2 thin and
seta-like, [Pmidapical spine or seta present]. Article
2 of pereopod 5 without facial setae. Peduncle of
uropod 1 with dorsolateral spines widely spread,
[Pmedial spines confined apically, Ppeduncle of uro-
pod 2 with only one medial spine or setule con-
fined apically]. Peduncle of uropod 3 lacking extra
subapical setae or spines. [PTelson with ordinary
pair of midlateral or dorsal setules on each side.]
TYPE-SPECIES.?Phoxus falcatus Sars, 1883 (mono-
typy). Monotypic, with 2 subspecies.
Leptophoxus falcatus falcatus (G. O. San)
Phoxus falcatus Sars, 1883:84.
Leptophoxus falcatus Sars, 1895:147-148, pi. 50.?J. L. Barn-
ard, 1960:308 [with references to include Phoxus simplex
Boeck, 1876:217-218. pi. 8: fig. 3].?Nagata, 1965:157.
DISTRIBUTION.?Greenland, east and west coasts;
Norway; North Sea; Skaggerak; Japan; 10-2258 m.
Leptophoxus falcatus icelus J. L. Barnard
Leptophoxus falcatus icelus J. L. Barnard, 1960:308-311, pis.
53, 54; 1966a:87; 1967:135.
DISTRIBUTION.?Southern California to middle
Baja California, 248-1120 m.
Metaphoxus Bonnier
Metaphoxus Bonnier 1896:630.?Stebbing 1906:138.?J. L.
Barnard 1960:303.
DIAGNOSIS.?Eyes present. Flagellum of antenna
2 reduced in female. Article 2 of antenna 1 short-
ened, ventral setae confined apically. Article 1 of
antenna 2 not ensiform; article $ with 2 setules;
facial spines on article 4 in 2 or more rows; article
5 ordinary in size to thin and short. Right man-
dibular incisor with 3 teeth; molar not triturative,
medium, elongate, conical, bearing one conical
fused spine, bearing fuzz; palpar hump small to
large. Palp of maxilla 1 uniarticulate; inner plate
naked. Setation of maxilla 2 weak. Inner plate of
maxillipeds ordinary; apex of palp article 3 not
protuberant, dactyl elongate to medium, apical nail
distinct, medium to elongate. Gnathopods similar
to dissimilar, enlarged, gnathopod 2 weakly to
strongly enlarged; article 5 gnathopods 1-2 very
short, cryptic, with eusirid attachment, palms
oblique to transverse to chelate, hands of gnatho-
pods 1-2 various, elongate or broadened, poorly
setose anteriorly. Article 5 of pereopods 1-2 setose
posteroproximally. Article 2 of pereopod 3 of
broad form, articles 4-5 of pereopods 3-4 narrow,
article 2 of pereopods 3-4 not setose posteriorly;
pereopod 5 ordinary, article 2 naked ventrally, arti-
cle 3 ordinary, dactyl normal. Epimera 1-2 lacking
long posterior setae, without midfacial setae above
ventral facial ridge; epimeron 3 of rounded classi-
fication and bearing 3 or fewer long setae. Uroso-
mite 1 generally naked except for sparse apicoven-
tral setae or spines near base of uropod 1; uroso-
mite 3 without dorsal hook or special process. Pe-
duncle of uropod I normally elongate, without
apicoventral spike, without special enlarged apico-
lateral-medial spine, peduncular apices of uropods
1-2 combed; inner ramus of uropod 1 with mar-
ginal spines in one row, no rami continuously spi-
nose to apex, apical nails weak to absent, inner
ramus of uropod 2 ordinary. Uropod 3 ordinary,
elongate article 2 of outer ramus carrying 2 short
to medium to vestigial apical setae. Telson elon-
gate, with 1-3 apical spines or setae on each lobe
plus setules, occasionally with supernumerary dor-
sal and lateral spines or setae.
DESCRIPTION.?Rostrum fully developed. Fuzz on
article 1 of antenna 1 in male present; calceoli on
male primary flagellum of antenna 1 present. Cal-
ceoli on article 5 of male antenna 2 present; fla-
gellum in male with calceoli. Prebuccal parts ordi-
nary, poorly separated from each other, upper lip
dominant. Right lacinia mobilis bifid, flabellate;
mandibular palp medium to thin, article 1 short,
article 2 without outer setae, apex of article 3
oblique. Lower lip bearing or lacking cones. Outer
plate of maxilla 1 with 7-9 spines, no spine espe-
cially thickened. Inner plates of maxilliped broad
to especially thin, poorly armed. Coxae 2-4 with-
out special anterodorsal humps. Some posterior
spines on article 6 of pereopods 1-2 thick and seta-
like, others thick and stiff, midapical spine or seta
present. Article 2 of pereopod 5 without facial
NUMBER 245 449
setae. Peduncle of uropod 1 with dorsolateral spines
confined apically, medial spines confined apically
or widely spread; peduncle of uropod 2 with only
one medial spine or setule confined apically. Pe-
duncle of uropod 3 lacking extra subapical setae
or spines. Telson with ordinary pair of midlateral
or dorsal setules on each side.
TYPE-SPECIES.?Metaphoxus typicus Bonnier 1896
(= Phoxocephalus pectinatus Walker, 1896) (mono-
py)
COMPOSITION.?Phoxocephalus fultoni Scott, 1890,
Metaphoxus frequens J. L. Barnard, 1960, and the
following new species: M. tuckatuck, M. mintus,
M. yaranellus, and M. tulearensis.
REMARKS.?Counts of facial spines on article 4
of antenna 2 appear unusual because the first row
at the corner often has medial members of suffi-
cient size or thickness to be counted as spines.
Key to the Species of Metaphoxus
(Compound key)
1. Palm of gnathopods 1-2 fully transverse, parachelate 2
Palm of gnathopods slightly to strongly oblique Choose 3 or 7
2. Pereopods 1-2 with stout locking spines M. tulearensis, new species
Pereopods 1-2 with thin setal locking "spines" M. fultoni
3. Outer ramus of uropod 1 distinct from other rami of uropods 1-2, either shorter, more
curved, or alone lacking apical nail 4
Outer ramus of uropod 1 like other rami of uropods 1-2 5
4. Inner ramus of uropod 1 naked, article 1 of outer ramus on uropod 3 naked laterally
M. tuckatuck, new species
Inner ramus of uropod 1 with one dorsal spine, article 1 of outer ramus on uropod 3 spinose
laterally M. mintus, new species
5. Article 6 of gnathopod 1 only 1.7 times as long as wide, edges not parallel, stenopodous part
of pereopod 5 elongate M. pectinatus
Article 6 of gnathopod 1 twice as long as wide, slender, edges parallel, stenopodous part of
pereopod 5 short 6
6. Article 6 of gnathopod 2 about 1.4 times as long as wide, telson with 2 apical spines on each
lobe, inner plate of maxilliped very thin, bearing one spine M. frequens
Article 6 of gnathopod 2 about twice as long as wide, telson with one apical spine on each
lobe, inner plate of maxilliped broad, bearing 4-5 spines M. yaranellus, new species
7. Article 6 of gnathopod 2 only 1.4 times as long as wide 8
Article 6 of gnathopod 2 about 1.8+ times as long as wide 9
8. Article 6 of gnathopod 1 twice as long as wide, slender, edges parallel, stenopodous part of
pereopod 5 short **? frequens
Article 6 of gnathopod 1 about 1.7 times as long as wide, edges usually not parallel, steno-
podous part of pereopod 5 elongate M. pectinatus
9. Outer ramus of uropod 3 naked laterally M. tuckatuck, new species
Outer ramus of uropod 3 spinose 1?
10. Article 6 of gnathopod 1 about 1.7 times as long as wide, dactyls of pereopods 1-2 smooth ..
M. mintus, new species
Article 6 of gnathopod 1 about 1.9 times as long as wide, dactyls of pereopods 1-2 with tiny
nodules M- yaroneUus, new species
Metaphoxus tuckatuck, new species
FIGURES 228-231
DESCRIPTION OF FEMALE (phenotype A).?Head
about 16 percent of total body length, greatest
width about 60 percent of length; rostrum reaching
apex of article 2 on antenna 1, unconstricted, ta-
pering evenly, apically rounded. Eyes large, with
purple core in preservative surrounded by large
ommatidia. Article 1 on peduncle of antenna 1
almost 1.4 times as long as wide, more than twice
as wide as article 2, apicoventral margin with sev-
eral setules, strongly produced dorsal apex with 3
setules; article 2 about 0.4 times as long as article
1, apicoventral corner with 2 setae, article 2 over-
riding article 3 weakly on medial surface; accessory
flagellum shorter than primary flagellum, 3-articu-
late; primary flagellum 5-articulate, aesthetascs
450
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 228.?Metaphoxus tuckatuck, new species, holotype, female "b," 3-46 mm, phenotype A
(a ?- male "a," 3.11 mm; c - male "c," 2.82 mm; u ? female "u," 3.05 mm).
NUMBER 245 451
present. Spine formula on article 4 of antenna 2 =
3-3-3, proximal set with one accessory setule, spines
thin, dorsal margin with 2 setae in tandem, ventral
face with 5 peak Mate setules, ventral margin with
about 6-7 short to long setae, apex with thin spine
and short seta; article 5 about 0.75 times as long as
article 4, facial spine formula = 2, with one acces-
sory setule, dorsodistal corner with 3 spine-setae,
dorsal margin smooth, ventral margin with 3 sin-
gle medium setae, ventrodistal apex with about 5
setae and setules; flagellum as long as article 4 of
peduncle, 4-articulate. Prebuccal complex convex
anteriorly, well extended forward, epistome and
upper lip mostly fused together, ventral margin of
upper lip weakly bisinuate. Palpar hump of man-
dible small; right incisor tridentate, with notch
denning second tooth and accessory hump between
teeth 2-3; left basically bidentate, distal branch
weakly bidentate and bearing weak proximal facial
hump; right lacinia mobilis flabellate, distal branch
poorly denned, margin toothed, face with weak
hump; left lacinia mobilis flabellate, 6-toothed:
right rakers 2, left 3; molar conical, setulose; palp
article 2 with subbasal seta and 4 subapical long
setae on inner margin, article 3 about as long as
article 2, oblique apex with 12 setae. Cones on
lower lip absent, mandibular lobes of medium
length, weakly attenuate and apically rounded, inner
lobes cleft about one third their length. Inner plate
of maxilla 1 large, ovate, naked; outer plate with 7
spines; palp exceeding apex of outer plate, bearing
2 apical and one weakly apicolateral setae. Inner
plate of maxilla 2 much broader and shorter than
outer plate, medial margin of inner plate with one
disjunct and distinctly medial seta plus 5 other
short to medium setae, outer plate with 2 apico-
lateral, 4 apical and one medial setae. Inner plate
of maxilliped with one mediofacial spine and 2
apical spines; outer plate with 5 spines, 2 longest
basal spines hidden behind inner plate; article 2
of palp with sparse, stout and short medial setae,
article 3 lacking facial setae, article 4 long, apical
spine of medium length, bearing 2 accessory set-
ules, first very long. Coxa 1 distally expanded and
broadly rounded anteroventrally; long setal for-
mula of coxae 1-4 = 8-9-9-3, each with tiny addi-
tional posteroventral bulbar setule; coxa 4 quad-
ratiform, posterodorsal excavation large to medium,
L-shaped, ratio of width to length =1:1 . Gnatho-
pod 2 scarcely stouter than gnathopod 1; long pos-
terior setae on article 2 of gnathopods 1-2 = 3-5
and 4; gnathopods subeusirid, article 5 small and
triangular but apex scarcely produced, article 6
almost slender, slightly more than twice as long as
wide, palm oblique, defining hump of medium size,
defining spine large, dactyl fitting palm, apex of
dactyl with flake, posterior margin of article 6
with 2-3 setae, posterior corner of article 5 almost
cryptic; length ratios of articles 5-6 on gnathopods
1-2 = 36:90 and 33:95, width ratios = 23:45 and
22:46. Pereopods 1-2 similar to each other; long
posterior setae on article 2 of pereopods 1-2 = 4
and 5-6; article 4 with about 4 apicoanterior setae
and 4 apicofacial setae on pereopod 1, 2 apicofacial
setae on pereopod 2, posterior margins of articles
4-5 moderately to weakly setose, article 5 with 1,
rarely 2 apicofacial setae, with long posterodistal
spine slightly exceeding apex of article 6, lacking
small contiguous spine at base (possibly represented
by facial seta), article 6 bearing one posterior seta,
2 apicofacial setae, one long apical spine, one short
spine and one seta, rows therefore composed of 2
and 2 with 2 accessory facial setae, lateral spine
row composed of shorter spine plus posterior seta,
(also interpreted as 3 + 2 + middle distal spine),
medial row composed of longer spine and one seta,
dactyl with apical flake and thin inner setule
(almost appressed), apical nail not evident. Coxae
5-7 posteroventral setule formula = 1-1-1. Articles
4-5 of pereopods 3-4 thin, facial spines absent,
pereopods 4-5 with one anterior facial ridge; width
ratios of articles 2, 4, 5, 6 of pereopod 3 =66:17:
15:7, of pereopod 4 = 80:19:13:8, of pereopod
5 = 92:13:11:6, length ratios of pereopod 3 =
78:24:24:28, of pereopod 4 = 95:52:47:45, of pereo-
pod 5=115:21:23:18; article 2 of pereopod 5
extending to apex of article 4, posterior margin
with small setule acclivities, ventral margin with
setule scallops; apex of article 6 finely combed.
Epimeron 1 with weak facial ridge, absent on epi-
meron 2, each with rounded-quadrate posteroven-
tral corner and convex posterior margin, each bear-
ing midsetule; epimeron 3 with weak posteroven-
tral tooth and sinus bearing setule, posterior mar-
gin deeply convex, bearing 2 setule notches, epi-
meron 2 with one facial seta, rarely with 2 in hori-
zontal tandem. Urosomite 1 ventrally naked, weakly
produced dorsally and bearing one setule. Peduncle
of uropod 1 with tiny basofacial seta, each apex
of peduncle with one medium spine, medial mar-
452 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 229.?Mctaphoxus tuckatuck, new species, holotype, female "b," 3.46 mm, phenotype A
(a ? male "a," 3.11 mm; c ? male "c," 2.82 mm).
NUMBER 245 453
gin with one spine, rami of uropods 1-2 naked
dorsally, inner rami of uropods 1-2 and outer ra-
mus of uropod 2 with apical flake, inner ramus of
uropod 1 with small mediobasal spinule, outer
ramus of uropod 1 slightly shortened, thinner and
more curved than inner ramus, uropod 2 with 2-3
dorsolateral spines on peduncle, medial apex with
one spine, apex of peduncle on uropods 1-2 with
lateral comb. Uropod 3 of medium length, pe-
duncle with 4 long ventrolateral spines, medially
with one short spine; rami feminine, inner ramus
extending to M. 100 on article 1 of outer ramus,
apex with immersed setule, article 2 of outer ramus
elongate, 0.50, bearing 2 apical setules, margins
naked, lateral apex of article 1 with 2 spines, me-
dial apex with one spine. Telson elongate, length-
width ratio about 16:11, apices tapering, narrow,
weakly excavate, each bearing medium spine and
small apicolateral setule, dorsal pair of plusetules
weakly diverse, near middle of each lobe, almost
directly centered. Bulbar setules of cuticle ex-
tremely sparse.
DESCRIPTION OF MALE (phenotype A).?Eyes
much larger than those of female, causing lateral
protrusion of head. Article 1 of antenna 1 with
medial patch of fuzz, disparity in length of flagella
greater than in female; primary flagellum bearing
calceolus (6-chambered) on each of articles 1-3.
Spine formula on article 4 of antenna 2 peduncle =
4?3-3, articles 3-4 with dorsomedial fuzz, article 5
with one dorsal calceolus and 2 groups of stiff male
setae, ventrodistal margin poorly setose; flagellar
formula = 24, 1-3, 5, 7 . . . 23. Apex of mandibu-
lar palp with 10 setae. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
2?(I?2)?2?3; article 6 of gnathopods slightly more
elongate than in female. Long setae on coxae
1-4 = (7-8)-8-(7-8)-3; coxa 4 smaller in relation
to coxae 1-3 than in female and very slightly nar-
rower than in female. Article 4 of pereopods 1-2
with 3?4 and 0 apicofacial setae. Article 2 of
pereopods 3?5 narrower than in female. Epimera
enlarged, epimeron 2 with 0-1 facial seta, epi-
meron 3 with 3 posterior setules besides one at
ventral corner. Urosome smaller than in female.
Apicomedial comer of peduncle on uropods 1-2
with pair of spines in largest male, only one spine
in smaller male. Rami of uropod 3 masculine,
inner reaching only to M. 90 on article 1 of outer
ramus, medial margins of rami setose. Telson
scarcely more elongate than in female, with basal
patches of dorsal denticles.
VARIATION (male phenotype A).?One outer plate
on maxilliped of male "a" bearing 4 spines of
stout kind.
ILLUSTRATIONS (phenotype A).?Specimens poorly
preserved, heads of males especially difficult to in-
terpret; dorsal view of male "c" head reconstructed
and one damaged eye omitted; anterior lobe of
coxa of pereopod 4 on male "c" omitted (damaged);
dactyls of pereopods 1-2 with apical flake pulled
away from dactyl in illustration, flake often closely
appressed to dactyl in many specimens; 2 facial
setae on epimeron 2 of male "a" represented only
by holes, setae broken away.
HOLOTYPE.?NMV, female "b," 3.46 mm, pheno-
type A.
TYPE-LOCALITY.?PPBES 919/4, 18 Nov 1971,
Port Phillip Bay, Victoria, Australia, 6 m, silty
sand.
VOUCHER MATERIAL (phenotype A).?Type-local-
ity: male "a," 3.11 mm (illus.); male "c," 2.82 mm
(illus.); female "u," 3.05 mm (illus.).
DIAGNOSIS OF PHENOTYPE B.?Ommatidia of ordi-
nary medium size; mandibular palp article 2 with
only 2-3 long inner apical setae; epimeron 2 with
1-4 setae; male right lacinia mobilis poorly serrate.
REMARKS.?This phenotype has the appearance
of a distinct species; our lack of extensive material
of juveniles in either phenotypes A or B prevents
us from tracing the life history of the two forms.
We would normally presume that phenotype A
represents terminal development and a pelagic-
neritic phase of this species as marked by the en-
larged ommatidia, but these specimens come only
from benthic grabs whereas phenotype B occurs
in neritic tows. Phenotype A usually has more than
twice as many setae on article 2 of the mandibular
palp as does phenotype B but usually has fewer
setae on epimeron 2, an asynchronous noncorrela-
tion between the two phenotypes suggesting that
they might be distinct species. The strange right
lacinia mobilis of the male of phenotype B is un-
explainable and also a possible mark of speciation.
If phenotype B develops into phenotype A then
the male right lacinia mobilis must revert to its
normal condition in terminal male phenotype A,
a strange situation. We know nothing of the life
history of the neritic phases of phoxocephalids but
suggest the possibility that phenotype B adults
454
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 230.?Metaphoxus tuckatuck, new species, holotype, female "b," 3.46 mm, phenotype A
(a ?? male "a," 3.11 mm; u ? female "u," 3.05 mm).
may also be terminal and that this species has, as
a survival mechanism, two terminal phases, A and
B.
DESCRIPTION OF PHENOTYPE B.?Like phenotype
A except in points of diagnosis.
OBSERVATIONS (phenotype B).?Female "m," 2.39
mm: Article 4 of antenna 2 with facial spine for-
mula of 3-3-2, basofacial setae 3 only, ventral mar-
gin with 4 long and one short setae only; article 5
with only 2 ventral setae. Mandibular palp article
2 with only 1-2 long inner distal setae, one short
basal seta, article 3 with only 9 apical setae. For-
mula of long setae on coxa 1-4 = 8-8-7-2. Epi-
meron 2 with 2 setae.
Female "e," size unknown: Ommatidia 11 in
number. Epimeron 2 with 2 setae.
Male "f," 3.22 mm: Giant specimen. Facial
spine formula on article 4 of antenna 2 = 2-2-2,
flagellar formula = 25, 1-4, 6, 8 . . . 24. Article 2
of mandibular palp with 4 long apical setae. Max-
illa 2 normal (like phenotype A; right mandible
illustrated). Setal formula of coxae l-^i = 6-7-6-2.
Epimeron 2 with 3 setae. Peduncle of uropod 1
with one apicolateral spine and one small spine in
middle of lateral margin. Uropod 2 with 4 spines
on peduncle.
Female "g," 2.71 mm: Ommatidia slightly en-
larged (illustrated).
Male "h," 2.70 mm: Palp article 2 of mandible
with 3 long inner distal setae. Formula of spines
on article 4 of antenna 2 = 1-3-2 only; article 5
like phenotype A but spines longer, article 4 with
NUMBER 245 455
wMt
FIGURE 231.?Metaphoxus tuck a tuck, new species, phenotype B (/ ? male "t," 3-22 mm; g ??
female "g," 2.71 mm; /i ? male "h," 2.70 mm; k ? male "k," 2.44 mm; m ? female "m," 239
mm; n ? female "n," 2.24 mm; w ? female "w," 2.41 mm; y ? male "y," 3.07 mm).
4 basofacial setae, article 5 with only 3 ventral
setules, one dorsal calceolus; flagellar formula =
20, 1-3, 5, 7 . . . 19. Formula of long setae on coxae
1-4 = 6-6-6-1. Article 4 of pereopods 1-2 with 2
and 0 facial setae. Article 2 of pereopod 4 slightly
stouter than in phenotype A. Epimeron 1 with only
one anterodorsal mediofacial seta; epimeron 2 also
with only one similar seta but with 4 lateral facial
setae. Peduncle of uropod 2 with only 2 lateral
peduncular spines and only one medial spine.
Inner ramus of uropod 3 with only 6 medial setae,
outer ramus with only 5 medial setae. Telson lack-
ing basal denticles.
Female "i," 2.5 mm: Like female "u" but
slightly larger. Ommatidia proliferating, each
doubled. Facial spine formula on article 4 of an-
tenna 2 = 3-3-3. Epimeron 2 with 2 setae. Apical
scales of uropods 1-2 especially attenuate and
elongate, peduncle of uropod 2 with 2 dorsal
spines.
Female "j," 3.04 mm: Like female "i" but om-
matidia not proliferating, total =14. Epimeron 2
with only one seta. Coxa 1 with 10 setae.
Male "k," 2.44 mm: Epimeron 2 with 3 lateral
facial setae. Uropod 2 with 3 lateral peduncular
spines; apices of rami on uropods 1-2 blunter than
in any other specimens of phenotypes A and B,
pointed flake poorly developed.
Female "w," 2.41 mm: Sexually mature but
very poorly developed, especially right lacinia
456 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
mobilis (illustrated). Primary flagellum of antenna
1 with 5 articles; accessory flagellum with S. Facial
spine formula on article 4 of antenna 2 = 3-2-2;
flagellum with 4 articles. Article 2 of right mandib-
ular palp with 3 long distal setae, left with 2. Long
setal formula of coxae 1-4 = 7-7-6-1. Epimeron 2
with 2 setae. Epimera 1, 3 and uropods 1, 2 normal.
Male "y," 3.07 mm: Eyes large but ommatidia
small. Antenna 2 with 5 basofacial plusetules on
article 4. Right lacinia mobilis somewhat interme-
diate between phenotypes A and B (illustrated).
Epimeron 2 with 3 setae. Peduncle of uropod 2
with 4 dorsal spines.
Male "z," 3.10 mm: Epimeron 2 with 3 setae.
Females associated with males "y" and "z":
Eyes small, ommatidia on stalks. Right lacinia
mobilis with strong, sharp teeth. Epimeron 2 with
2 setae.
VOUCHER MATERIAL (phenotype B).?RHM:
male "f," 3.22 mm (illus.); female "g," 2.71 mm
(illus.); male "h," 2.70 mm (illus.); male "k," 2.44
mm (illus.); female "m," 2.39 mm (illus.); female
"n," 2.24 mm (illus.); males "p," "q," "s," juvenile
"t." CPBS 000/5: female "w," 2.41 mm (illus.).
PPBES 906/4: male "y," 3.07 mm (illus.); male "z,"
3.10 mm. CPBS 03S/2: female "e," size unknown.
CPBS M00/1: female "i," 2.5 mm. CPBS 100/5:
female "j," 3.04 mm.
RELATIONSHIP.?This species is generally similar
to M. frequens J. L. Barnard, I960, from California
but the latter differs from M. tuckatuck in the fol-
lowing characters: the stouter article 6 of gnatho-
pod 2, the presence of only one spine on the inner
plate of the maxilliped, the absence of setules on
epimeron 3, the presence of one dorsal spine on the
outer rami of uropods 1-2, the less strongly setose
maxilla 2, fewer coxal setae, small ommatidia,
fewer setae on article 2 of mandibular palp, poorly
developed accessory setule on article 4 of maxil-
lipedal palp, presence of 2 setae on the inner plate
of maxilla 1 and the presence of 2 spines and one
setule on each apex of the telsonic lobes.
MATERIAL.?CPBS, 16 samples from 11 stations
(19); WPBES, 7 samples from 5 stations (15);
PPBES, 79 samples from 41 stations (143); RHM,
one sample {10).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 0-24 m and neritic, sand, muddy sand,
sand and gravel, clay, silty clay, often heavy sea-
grass cover.
Metaphoxus yaranellus, new species
FICHUS 232. 233
DESCRIPTION OF MALE.?Head about 17 percent
of total body length, greatest width about 66 per-
cent of length; rostrum reaching middle of article
2 on antenna 1, unconstricted, tapering evenly,
apically rounded. Eyes medium to medium-large,
eosin \i\ color, clear of occluding pigment, om-
matidia ovate or elliptical in small-eyed males, be-
coming elongate in terminal males. Article 1 on
peduncle of antenna 1 about 1.2 times as long as
wide, twice as wide as article 2, apicoventral
margin poorly setulose, produced dorsal apex with
one setule, medial face densely fu//y. lutz appar-
ently composed of thick aesthetascs; article 2 about
0.65 times as long as article 1, apicoventral lateral
face with 4-5 setae, article 2 slightly overriding
article 3 on medial surface; accessory flagellum
slightly shorter than primary flagellum, 3-articu-
late; primary flagellum 4-articulate, bearing one
calceolus each on articles 1 or 1-2 and several
aesthetascs. Spine formula on article 4 of antenna
2 = 1-2-2-3 or 1-2-2-4, set 3 with one accessory
setule, set 4 with 1-3 accessory setules, spines thin,
dorsomedial margin densely fuzzy, fuzz composed
of thick aesthetascs, proximoventral margin with
2-3 penicillate setules, distally with pair of setae,
ventral apex, with spine and 3 setae; article 5
about 0.8 times as long as article 4, facial spine
formula = 2, no accessory setule, dorsodistal corner
with 2 spine-setae, dorsal margin with 0-1 calceolus
and 1-2 groups of male setae, ventrodistal apex
with one spine-seta, pair of short setule-spinules,
ventral margin with pair of setules; flagellum
elongate, flagellar formula = (15-17), 1 ,2 ,4 ,6 ,8
only or 1, 3, 5 only or 1, 3, 5 . . . 15 (from 9 onward
calceoli rudimentary), or 1, 2, 3, 5 only, or article 2
only. Prebuccal complex convex anteriorly, poorly
extended forward, epistome and upper lip mostly
fused together, ventral margin of upper lip weakly
bisinuate. Palpar hump of mandible medium,
broad; right incisor basically tridentate, with
fourth tooth adjacent to tooth 2; left basically bi-
dentate, distal branch strongly tri- or quadriden-
tate; right lacinia mobilis flabellate, proximal
branch distinct from serrate distal branch, face
naked; left lacinia mobilis flabellate, 5-toothed;
right rakers 3 plus 1-3 rudimentaries; left rakers 3
plus one rudimentary; molar conical, setulose; palp
NUMBER 245 457
FIGURE 232.?Metaphoxus yaranellus, new species, holotype, male "a," 234 mm (n ? male "n,"
2.63 mm).
458 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
article 2 with inner subbasal setule and 2-3 sub-
apical long setae, article 3 about 0.65 times as long
as article 2, oblique apex with 8 setae. Each outer
lobe of lower lip with cone, mandibular lobes long,
weakly attenuate and apically rounded, inner lobes
cleft about one-third. Inner plate of maxilla 1 of
medium size, ovate, bearing one apicomedial
setule; outer plate with 7 spines; palp slightly ex-
ceeding apex of outer plate, bearing 2 apical and
one apicolateral setae, occasionally a fourth apico-
medial seta. Inner plate of maxilla 2 slightly
broader and much shorter than outer plate; medial
margin of inner plate with one disjunct medial
seta plus 5-7 other apicad setae of which one very
stout; outer plate with 2 apicolateral, 6 apical and
2 medial setae. Inner plate of maxilliped with one
mediofacial spine, one apicomedial spine, and 2-3
apical spines; outer plate with 3 apicomedial
spines, one elongate basomedial spine; article 2 of
palp with sparse, stout medial setal-spines, article
3 lacking facial setae, article 4 long, apical spine of
medium length, bearing 2 accessory setules, first
setule elongate. Coxa 1 scarcely expanded; long
setal formula of coxae 1-4 = (2-5)-(4-5)-(3-5)-(0),
each with tiny additional posteroventral setule;
coxa 4 quadratiform, posterodorsal excavation
medium-small, V-shaped, posterodorsal corner
rounded, ratio of width to length = 7:9. Gnatho-
pod 2 stouter and longer than gnathopod 1; long
posterior setae on article 2 of gnathopods 1-2 = 1
and 1; article 5 small and triangular; article 6
elongate, twice as long as wide, palm weakly
oblique, defining hump medium to large, defining
spine large, dactyl fitting palm, apex of dactyl with
flake, posterior margin of article 6 with 1-2 setae;
length ratios of articles 5-6 on gnathopods 1-2 =
48:87 and 46:98, width ratios = 24:47 and 26:54.
Pereopods 1-2 similar to each other; long pos-
terior setae on article 2 of pereopods 1-2 = 2
and 2; article 4 with one long apicoanterior seta,
no apicofacial setae, posterior margin with sets of
3 and 2 setae on pereopod 1, 4 and 3 on pereopod
2; article 5 with one apicofacial seta, long postero-
distal spine reaching M. 80 on article 6, with small
contiguous spinule at base; article 6 bearing one
posterior seta and apical sets of 3 and 2 spines of
medium length; dactyl with apical immersed nail,
appearing to have apicofacial meat us, thin inner
setule on weak acclivity, inner margin and lateral
face bearing minute rows of nodules. Coxae 5-7
posteroventral setule formula = 1?4?4. Articles 4-5
of pereopods 3-4 thin, facial spines absent, pereo-
pods 4-5 with one anterior facial ridge; width
ratios of articles 2,4,5,6 of pereopod S =
48:24:18:8, of pereopod 4 - 7225:179, of pereo-
pod 5 = 110:20:157, length ratios of pereopod
3 = 68:27:29:32, of pereopod 4 = 91:49:46:42. of
pereopod 5 = 128:28:28:16; article 2 of pereopod
5 especially large, reaching middle of article 5,
posterior margin with small setule acclivities, ven-
tral margin with weak setule scallops; apex of
article 6 finely combed. Epimeron 1 with facial
ridge, absent on epimeron 2 (illustration showing
ventral margin of next instar), each with rounded
posteroventral corner and weakly convex posterior
margin bearing 2 setules, epimeron 3 with deeply
convex and notched posterior margin, posteroven-
tral corner with *t?tule notch, epimeron 2 with
2-4 facial setae in horizontal tandem. Urosomite 1
bearing one setal spine midventrally on each side.
Peduncle of uropod 1 naked basofacially, lateral
apex with one medium spine, medial apex with
longer spine, medial margin with 1-2 additional
spines, rami of uropods 1-2 naked dorsally, apices
grossly naked and apparently lacking immersed
nail (probably fully absorbed), inner rami often
with apical flake, outer rami rarely so, inner
ramus of uropod 1 with small mediobasal setule-
spinule, rami of uropod 1 extending equally,
peduncle of uropod 2 with 2 dorsal spines, medial
apex with one spine, apex of peduncle on uropods
1-2 with lateral comb. Uropod 3 elongate, inner
ramus strongly setose medially but not fully elon-
gate (not fully adult), peduncle with 4-5 ventro-
lateral spines, medially with short spine and longer
seta, dorsolaterally with setule, inner ramus extend-
ing to M. 75 on article 1 of outer ramus, apex with
one long seta, one setule, article 2 of outer ramus
elongate, 0.50, bearing 2 minute apical setules, lat-
eral margin of article 1 with 3 acclivities; seta for-
mula = l(short)-l-l-l, spine formula = 0?1-1-1
(variable, setae = 1-1-1-0, spines = 0-1-1-2, or
setae = 1-1-1-1 and spines = 0-1-1-2), medial
apex with one spine, one seta. Telson highly elon-
gae, length-width ratio about 7:4, apices tapering,
narrow, apically truncate, bearing one short spine
in middle of apex and one apicomedial setule,
dorsal pair of plusetules alike, near middle of each
lobe close to base, dorsal cuticle of telson covered
with fuzz except for basal triangle bearing chisel
NUMBER 245 459
E2
FIGURE 233.?Metaphoxus yaranellus, new species, holotype, male "a," 2.54 mm (n - male "n,
2.63 mm).
460 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY
shaped denticles in subcircular group, other stout
setules distally. Cuticle covered with fuzz easily
visible under medium and low power microscopy
and scattered bulbar setules.
VARIATIONS.?Apical flakes on rami of uropods
1-2 variable in occurrence, usually borne only on
inner rami, often absent, very rarely on outer rami;
male "b" with apicomedial spine position on
peduncle of uropod 1 bearing pair of spines; only
males "a" (holotype) and "d" with oddly elongate
ommatidia.
ILLUSTRATIONS.?Head of holotype slightly flat-
tened laterally; holotype telson shown flattened,
telson of male "n" shown flattened; body and ap-
pendages including uropods and telson almost fully
covered with cuticular fuzz but, in those illustra-
tions showing fuzz, only highlights actually de-
picted, except for naked triangle on base of telson.
HOLOTYPE.?AM, male "a," 2.54 mm.
TYPE-LOCALITY.?AM P. 18125, 12 Dec 1939,
Antechamber Bay, Kangaroo Island, South Aus-
tralia, with light in net at night.
VOUCHER MATERIAL.?Type-locality: male "b,"
2.61 mm; male "n," 2.63 mm (illus.); male "d," 2.40
mm; and male "e," 2.41 mm. Female unknown.
RELATIONSHIP.?This species differs from Mela-
phoxus tuckatuck in the unshortened outer ramus
of uropod 1, the very narrow coxa 4, fuzzy cuticle,
enlarged article 2 of pereopod 5, elongate telson
with apical setule on medial side of main spine,
the greater spinosity on the lateral face of article 4
on antenna 2, the poorly expanded coxa 1 and the
shapes of gnathopods 1-2.
Metaphoxus yaranellus bears the vaguest resem-
blance to M. frequens J. L. Barnard, 1960, from
the northern hemisphere, but differs in numerous
characters involving gnathopod 2, epimeron 3,
mouthparts, telson, coxae, uropods, and pereopod
5.
MATERIAL.?AM, one sample (5).
DISTRIBUTION.?South Australia, Kangaroo Island,
neritic.
Metaphoxus tnintus, new species
FIGURES 234-236
DESCRIPTION OF FEMALE.?Head about 17 per-
cent of total body length, greatest width about 55
percent of length; rostrum exceeding apex of arti-
cle 3 on antenna 1, unconstricted, narrow, elongate.
Eyes large, largely occluded with pigment, om-
matidia small. Article 1 on peduncle of antenna 1
about 1.4 times as long as wide, about 1.8 times
as wide as article 2, apicoventral margin and face
with about 9 setules, strongly produced dorsal apex
with 3 setules; article 2 about 0.15 times as long
as article 1, with apicoventral cycle of 4-5 setae,
article 2 scarcely overriding article 3 on medial
surface; accessory flagellum shorter than primary
flagellum, 3-articulate; primary flagellum 6-articu-
late. Spine formula on article 4 of antenna 2 =
3-3-5 or 3-3-4, set 2 with one accessory setule, set
3 with 5 accessory setules, spines thin, dorsal
margin with 3 sets of 1-3 setae, ventral margin
with 6-7 sets of 1-3 long to medium setae, ventral
apex with one medium spine; article 5 about 0.63
times as long as article 4, facial spine formula -
3-2, each group with accessory seiulc, dorsodistal
corner with 2 large spines, dorsal margin naked,
ventral margin with 2 sets of 2 long to medium
setae, ventrodistal apex with 3 long to medium
spine-setae; flagellum about 0.63 times as long as
articles 4-5 of peduncle combined, with 6 articles.
Prebuccal mass flat, weakly extended anteriorly,
ventral margin of upper lip with small hump.
Palpar hump of mandible medium-strong; right
incisor basically tridentate, with fourth tooth ad-
jacent to tooth 2; left basically bidentate, distal
branch bifid; right lacinia mobilis flabellate, distal
branch distinct from simple proximal branch, face
naked; left lacinia mobilis flabellate, 5-toothed,
with accessory tooth; right rakers 2, left 3; molar
subconical, setulose; palp article 2 with 3 inner
subbasal setae and 5 longer subapical setae in
tandem, article 3 about 0.9 times as long as article
2, oblique apex with 13-14 setae, basofacial for-
mula = 0-1. Each outer lobe of lower lip with
cone, mandibular lobes short, weakly attenuate and
apically rounded, inner lobes cleft about one third.
Inner plate of maxilla 1 of medium size, ovate,
naked; outer plate with 7 spines; palp slightly ex-
ceeding apex of outer plate, bearing 3 apical and
one apicolateral setae. Inner plate of maxilla 2
slightly broader and much shorter than outer plate,
medial margin of inner plate with 5-6 setae, outer
plate with 2-3 apicolateral, 13-14 apical and 2
medial setae. Inner plate of maxilliped with 2
mediofacial setae, one apicomedial spine, 4?5
apical spines, outer plate with 6 apicomedial
NUMBER 245
\o Jf ^ -?^^
s
FIGURE 234.?Metaphoxus mintus, new species, holotype, female "a," 4.95 mm (n = male "n,"
4.70 mm).
462 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY
spines, one elongate basomedial spine, article 2 of
palp with dense medial setation, article 3 with 2
facial setae, article 4 long, bearing 2 accessory
setules, apical spine short. Coxa 1 expanded dis-
tally; long setal formula of coxae 1-4 = (11-15)-
(8-12)-(7-9)-(2-4), some middle setae shortened,
each with tiny additional posteroventral setule;
coxa 4 quadratiform, anterior and posterior mar-
gins weakly divergent, posterodorsal excavation
medium, V-shaped, posterodorsal corner rounded,
ratio of width to length = 17:19. Gnathopod 2
scarcely stouter but longer than gnathopod 1; long
posterior setae on article 2 of gnathopods 1-2 = 3
and 2-3; article 5 small, triangular; article 6
elongate, almost twice as long as wide, palm
oblique, defining hump small, defining spine med-
ium, dactyl overriding palm, apex of dactyl with
flake, posterior margin of article 6 with 3 setae;
length ratios of articles 5-6 on gnathopods 1-2 =
40:82 and 44:89, width ratios = 23:48 and 25:48.
Pereopods 1-2 similar to each other; long posterior
setae on, article 2 of pereopods 1-2 = 3 and 4;
articles 4-5 each with apicofacial row of 3 setae,
posterior margin of article 4 with 4 sets of 3-7
setae, long posterodistal spine on article 5 reaching
M. 95 on article 6, with tiny contiguous spinule at
base; article 6 bearing 2 posterior setae and apical
sets of 2 and 2 spines, mostly thin, plus middistal
enlarged spine-seta; dactyl with apical immersed
nail, thin inner setule on weak acclivity, surface
naked. Coxae 5-7 posteroventral setule formula =
1-1-1. Articles 4-5 of pereopods 3-4 thin, facial
spines absent, pereopods 4-5 with one anterior
facial ridge; width ratios of articles 2, 4, 5, 6
of pereopod 3 = 60:20:17:8, of pereopod 4 =
65:22:16:8, of pereopod 5 = 86:19:16:8, length
ratios of pereopod 3 = 80:28:27:27, of pereopod
4 = 89:44:38:34, of pereopod 5=118:28:26:16;
article 2 of pereopod 5 especially large, reach-
ing apex of article 4, posterior margin with
weak setule acclivities, ventral margin with
weak setule scallops; apex of article 6 finely
combed. Epimeron 1 with facial ridge, absent on
epimeron 2, each epimeron with rounded postero-
ventral corner and straight to weakly convex pos-
terior margin bearing one setule, epimeron 3 with
weakly protuberant and 2-notch posterior margin,
posteroventral corner with notch, epimeron 2 with
4-8 facial setae. Urosomite 1 naked. Peduncle of
uropod 1 naked basofacially, lateral apex with one
medium spine, medial apex with one spine, rami
of uropods 1-2 naked dorsally except for one spine
on inner ramus of uropod 1, apices of all but outer
ramus on uropod 1 with scarcely distinct tiny
apical nail, inner ramus of uropod 1 with small
mediobasal setule-spinule, outer ramus of uropod 1
slightly shorter and more curved than inner ramus,
peduncle of uropod 2 with 1-2 dorsal spines,
medial apex with one spine, apex of peduncle on
uropods 1-2 with lateral comb. Uropod 3 weakly
elongate, peduncle with 5 ventrolateral spines,
medially with 2 spines, dorsolaterally with one
spine; inner ramus extending to M. 85 on article 1
of outer ramus, apex with 2 long setae, otherwise
naked, article 2 of outer ramus elongate, 0.40, bear-
ing 2 short setules, lateral margin of article 1 bear-
ing 2 acclivities; setal formula = 0, spine formula
= 1-1-2, medial apex with pair of setal-spines.
Telson highly elongate, length-width ratio about
17:11, not fully cleft, apices tapering, narrow,
weakly rounded or note lied, bearing one short
spine in middle of apex, one apirolateral setule,
often one medial setule, dorsal pair of plusetules
weakly diverse, near middle of lobe, base with row
of denticles on each lobe. Cuticle with sparse
bulbar setules, surface bearing patches of fine stria-
tions in form of fingerprint pattern, emergent
setules especially short.
VARIATIONS (female).?Female "b" with 6 facial
setae on epimeron 2, female "w" with 8, inner
ramus on uropod 1 of female "b" with 2 dorsal
spines, but both females "b" and "w" with only
3-0 spines on article 5 of antenna 2; female "w"
with aberrant thin spine on dorsobasal face of
inner ramus on uropod 1, epimeron 1 with one
anterofacial seta, apex of peduncle on uropod 2
with dorsal half of comb much longer and com-
posed of stouter elements than in other specimens
except juvenile "g," thus female "w" bearing ap-
parent juvenile condition on uropod 2, though gen-
erally more senile than female "a" (holotype) and
"b."
DESCRIPTION OF MALE (male "n," 4.70 mm).?
Eyes scarcely larger than in female, some om-
matidia larger. Article 1 of antenna 1 with medial
patch of fuzz, articles 1-4 of primary flagellum
each with calceolus, terminal articles with multi-
ples of elongate aesthetascs. Dorsomedial faces of
articles 3-4 of antenna 2 fuzzy; spine formula on
face of article 4 = 3-3-4, ventral setae short but
NUMBER 245 463
V)P5
FIGURE 2S5?Mctaphoxus mintus, new species, holotype, female "a," 4.95 mm (a; ? female "w,1
4.51 mm; x = left).
not reduced in number; article 5 enlarged,
ventrally and distally with setae reduced in number
and shortened, facial spine formula = 2-0, anterior
margin with one calceolus and bundles of male
setae; flagellum elongate, flagellar formula = 32,
1-3,5,7 . . . 31. Upper lip almost evenly truncate
ventrally. Right lacinia mobilis almost smooth, not
bifid, forming asymmetrical arrow-head. Palpar
apex of maxilla 1 with 5 setae. Apex on inner plate
of maxilliped with only 3 of stout kind of spine,
outer plate with only 5 thick and one thin spines.
Long setae on coxae 1 ^ = (11-12)-(8)-(6-7)-(2),
coxa 1 with one long seta anterior to anteroventral
bulbar setule(I); coxa 4 slightly larger relative to
coxa 1 than in female, otherwise narrower. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 3-1-3-4, short anterior setae =
1-5-3-3, no others. Lateral facial setae on articles
4-5 of pereopods 1-2 = 2-2 and 0-2; article 6 with
spine rows of 2 + 2 plus middistal seta plus one
posterior seta. Article 2 of pereopod 5 scarcely nar-
rower than in female but entire appendage larger
relative to size of pereopod 3 than in female. Epi-
mera enlarged, epimeron 2 with 6 facial setae,
epimeron 3 with 3 posterior setules above corner
setule. Uropod 3 masculine, inner ramus almost
extending to apex of article 1 on outer ramus,
setose medially and laterally, article 1 of outer
464 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 236.?Metaphoxus mintus, new species, holotype, female "a," 4.95 mm (n ? male "n,f
4.70 mm).
ramus with 2 lateral acclivities but spine formula
= 0-0-1-1-2 and setal formula = 1-1-1-1-1.
Telson even longer than in female, each apex with
additional apical setule medial to main spine, lat-
ter shortened, basodorsal denticles more numerous
than in female.
OBSERVATIONS.?Male "f," 4.48 mm: Flagellar
formula on antenna 2 = 27, 1, 2,4, 6 . . . 26. Long
setae on coxae 1-4 = 12-9-8-2. Epimeron 2 with 5
setae. Article 1 on outer ramus of uropod 3 with 3
acclivities, spine formula = 0-1-1-2, setal formula
= 1-1-1-1.
DESCRIPTION OF JUVENILE.?Juvenile "g," 3.00
mm: Eyes of medium size. Article 2 of antenna 1
with 2 ventrodistal setae; primary flagellum with 5
articles; accessory flagellum with 3 articles. Spine
formula on article 4 of antenna 2 = 3-3-2, on arti-
cle 5 = 2-0; flagellum with 4 articles. Long setae
NUMBER 245 465
on coxae 1,2, and 4 = 8-6-0. Epimeron 2 with 3
facial setae. Peduncle of uropod 2 with one dorsal
spine, upper half of apical comb enlarged, elements
longer and thicker than in ordinary adults (but see
female "w"). Article 1 on outer ramus of uropod 3
with one lateral acclivity, spine formula = 1-2,
setal formula = 0-0, apex of inner ramus with one
setal spine.
Juvenile "h," 3.10 mm: Accessory flagellum of
antenna 1 3-articulate; primary flagellum 5-articu-
late. Spine formula on article 4 of antenna 2 =
3-3-3, on article 5 = 3-0. Right lacinia mobilis
with distal flabellate branch bearing 6 teeth
divided into 2 groups of 3 each separated by sinus.
Apex of palp on maxilla 1 with 3 distal setae and
one apicolateral seta. Long setae on coxae 1-4 =
11-6-5-1. Epimeron 2 with 5 setae. Upper half of
comb on peduncle of uropod 2 enlarged. Inner
ramus of uropod 3 with one apical spine-seta, outer
ramus with one lateral acclivity, spine formula =
1-2, setal formula = 0-0.
ILLUSTRATIONS.?Outer ramus on female uropod
3 disarticulate in drawing.
HOLOTYPE.?AM, female "a," 4.95 mm.
TYPE LOCALITY.?SBS C1S5, 17 May 1972, off
Malabar, New South Wales, Australia, 29 m, shelly
gravel, high organic content.
VOUCHER MATERIAL.?Type-locality: female "b,"
4.82 mm; female "w," 4.51 mm (illus.); male "n,"
4.70 mm (illus.); male "f," 4.48 mm; juvenile "g,"
3.00 mm. SBS A2S5: juvenile "h," 3.10 mm.
RELATIONSHIP.?The slightly shortened and
curved outer ramus of uropod 1 suggests relation-
ship to Metaphoxus tuckatuck.
The strongest clue to the distinction of Af.
mintus is the blunt apices on the rami of uropods
1-2 but the following characters of M. mintus also
appear to confirm its identity: inner ramus of uro-
pod 1 with dorsal spine; presence of 2 long and
thick apical setae on inner ramus of uropod 3;
stronger spinosity and setosity of epimeron 2 and
article 4 of antenna 2; stronger distal armament on
the inner plate of the maxilliped; presence of lat-
eral spines and setae on article 1 of the outer ramus
on uropod 3; posteriorly truncate coxa 5; stouter
and distinctly shaped articles 4-5 of pereopod 5;
and shapes of right lacinia mobilis in both sexes.
MATERIAL.?SBS, 7 samples from 5 stations (16,
including 2 dubious).
DISTRIBUTION.?New South Wales, off Malabar,
29-48 m, shelly gravel and medium fine sand.
Metaphoxus frequens J. L. Barnard
Metaphoxus frequens J. L. Barnard, 1960:504-306, pl$. 51, 52;
1964c:242; 1966a:88; 1966b:28; 1969b: 196; 1971:68.
DISTRIBUTION.?Oregon to Isla Isabel, Mexico,
13-458 m.
Metaphoxus fultoni (Scott)
Phoxocephalus Fultoni Scott, 1890:327-328; pi. 12: figs. 10-
12, pi. 13: figs. 13-19; 1906:148.?Chevreux, 1898:477 ?
Norman, 1900:336.
Phoxocephalus chelatus Delia Valle, 1893:742-743, pi. 5: fig.
10, pi. 35: figs. 29-35.
Phoxocephalus fultoni.?Robertson, 1892:207-208.?Walker,
1895:296.?Caiman, 1896:743-748, pi. 31: figs. 1-2.?Scott,
1907:178.?Patience, 1909:118-125, pis. 3, 4 [part].
Metaphoxus Fultoni.?Walker, 1901:299? Norman and Brady
1909:306.?Chevreux, 1911:189, pi. 9: figs. 17, 18.?Chevreux
and Fage. 1925:106-107, figs. 96, 100.?Fage, 1933:203-207.
?Candeias, 1934:2.
Metaphoxus fultoni.?Stebbing, 1906:139.?Tattersall, 1913:
6-7.?Stephensen, 1929:84, fig. 21:99.?Plymouth Marine
Fauna 1931:189.-Moore, 1937:119.?J. L. Barnard, 1960:
304; 1964a: 103. fig. 18; 1964c:242-243; 1966b:28; 1969b: 196.
?Hamond, 1967:119.?Ledoyer, 1968:191.?G. S. Karaman,
1973:79-34, figs. 18-20.
DISTRIBUTION.?Eastern Atlantic and Mediterran-
ean from Adriatic Sea westward and northward to
British Isles; eastern Pacific Ocean from Monterey
Bay south to Isla Isabel, Mexico, 0-458 m.
Metaphoxus pectinatus (Walker)
Phoxus simplex Bate, 1857:525 [fide Norman, 1900, not Walk-
er, 1896b]; 1862:97-98, pi. 16: fig. 1.?Bate and Westwood,
1863:140-142, fig. [unnumbered].?Boeck, 1871:135.?Rob-
ertson, 1888:27-28.
Phoxus Kroyerii Bate, 1857:140 [= homonym, not Stimpson,
1853].
Phoxocephalus simplex.?Caiman, 1896:748-751, pi. 32: fig. 3.
?Norman, 1900:335-336.
Phoxocephalus pectinatus Walker, 1896a: 343-344, pi. 16: figs.
1-6; 1896b: 156-157.
Metaphoxus typicus Bonnier, 1896:630-633, pi. 37: fig. 1 ?
Chevreux, 1927:72.
Metaphoxus fultoni?Patience, 1909:125-130, pis. 3, 4: fig. 1
[in part, not Scott, 1890].
Metaphoxus pectinatus?Chevreux, 1898:477; 1900:33-34 ?
Walker, 1901:299-300, pi. 27: fig. 22.?Stebbing, 1906:139-
140.?Chevreux, 1911:187-188, pi. 9: figs. 19-20; 1925:295;
1935:73.?Tattersall, 1913:7.?Chevreux and Fage, 1925:
466 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
105-106, figs. 98, 99.?Plymouth Marine Fauna, 1931:189.?
Fage, 1933:205-207, fig. 10?Ruffo, 1938:130; 1947:84 ?
Soika, 1949:197?J. L. Barnard. 1960:307-308.?Bcllan-
Santini, 1965:4.?Ledoyer, 1968:191; 1970:15.?G. S. Kara-
man, 1973:84-91, figs. 21-24.
DISTRIBUTION.?Eastern Atlantic Ocean and
Mediterranean Sea, from the Adriatic Sea westward
and northward to the British Isles, meropelagic
and benthic to 80 m.
Metaphoxus tulearensis, new species
Metaphoxus fultoni.?Ledoyei, 1967:28, pi. 7 [not Scott].
DIAGNOSIS.?Like Metaphoxus fultoni (Scott) but
pereopods 1-2 with pair of stout locking spines at
base of dactyl; gnathopod 1 parachelate.
HOLOTYPE.?Female, 4 mm, in plate 7 of
Ledoyer, 1967.
TYPE-LOCALITY.?Station 106, Madagascar.
DISTRIBUTION.?Madagascar.
Metaphoxoides Ledoyer
Metaphoxoides Ledoyer, 1970:29-30.
DIAGNOSIS.?Eyes present. [Flagella of antennae
1-2 unreduced in female in M. zavorus, unknown
in type.] Article 2 of antenna 1 shortened to ordi-
nary, ventral setae confined apically. [PArticle 1 of
antenna 2 not ensiform; article 3 with 4 setae in
M. zavorus, unknown in type;] facial spines on
article 4 in 2 rows; article 5 ordinary (in type) to
especially thin; right mandibular [incisor un-
known]; molar not triturative [possibly absent];
palpar hump large. Palp of maxilla 1 uniarticulate;
inner plate naked (type) or with 3 setae. Setation
of maxilla 2 weak. Inner plate of maxillipeds ordi-
nary; apex of palp article 3 not protuberant, dactyl
elongate, apical nail distinct, elongate. Gnathopods
dissimilar, enlarged; article 5 of gnathopod 2 very
short, cryptic, ordinary on gnathopod 1, without
eusirid attachment, palms transverse to oblique re-
spectively, hands of gnathopods broadened, poorly
setose anteriorly. [Article 5 of pereopods 1-2 un-
known.] Article 2 of pereopod 3 of broad form,
articles 4-5 of pereopods 3-4 broad to narrow, arti-
cle 2 of pereopods 3-4 not setose posteriorly; pereo-
pod 5 ordinary, article 2 naked ventrally, article
3 ordinary, dactyl normal. [PEpimera 1-2 lacking
long posterior setae, without midfacial setae above
ventral facial ridge]; epimeron 3 ordinary, of
rounded classification and lacking long setae.
[PUrosomite 1 generally naked; urosomite 3 with-
out dorsal hook or special process.) Peduncle of
uropod 1 normally elongate, without apicoventral
spike, without special enlarged apicolatetal-medial
spine, [Ppeduncular apices of uropods 1-2 not
combed], inner ramus of uropod 1 lacking mar-
ginal spines, no rami continuously spinose to apex,
inner ramus of uropod 2 ordinary. Uropod 3 ordi-
nary, outer ramus with or without second article
bearing 2-4 long apical setae. Telson ordinary,
with 1-4 apical spines or setae on each lobe.
DESCRIPTION.?Rostrum fully developed. [Fuzz on
article 1 of antenna 1 in male unknown. Calceoli
on male antennae 1-2 unknown. PPrcbuccal parts
ordinary, poorly separated from each other, upper
lip dominant. Right lacinia mobilis Uabellate;]
mandibular palp medium to thin, article 1 short to
slightly elongate, article 2 without outer setae, apex
of article 3 oblique or rounded (type). [?Lower lip
lacking cones.] Outer plate of maxilla 1 with 7
spines, no spine especially thickened. Inner plates
of maxilliped especially thin, poorly armed.
[PCoxae 2-4 without special anterodorsal humps.
PSome posterior spines on article 6 of pereopods
1-2 thin and seta-like, others thick and stiff, Pmid-
apical spine or seta present.] Article 2 of pereopod
5 without facial setae. Peduncle of uropod 1 with
dorsolateral spines confined apically, medial spines
confined apically; [Ppeduncle of uropod 2 with only
one medial spine or setule confined apically].
Peduncle of uropod 3 lacking extra subapical setae
or spines. [PTelson with ordinary pair of midlateral
or dorsal setules on each side, Preduced to one
member on each side.]
TYPE-SPECIES.?Metaphoxoides picardi Ledoyer,
1970 (monotypy).
COMPOSITION.?Metaphoxus zavorus, new species
(= Metaphoxus sp., Griffiths, 1973).
RELATIONSHIP.?The vestigiality of article 2 on
the outer ramus of uropod 3 is the primary syste-
matic character of this genus.
Metaphoxoides zavorus is placed provisionally in
this genus until more facts can be ascertained about
the species. It was based on one broken specimen.
The vestigial article 2 of the outer ramus on uro-
pod 3 is a clue to its relationship, and there are
numerous other similarities between M. zavorus
NUMBER 245 467
and Af. picardi. However, Af. zavorus differs in
these potentially crucial characters: poorly devel-
oped nail on dactyl of maxilliped in contrast to a
well developed nail with large accessory nails on
Af. picardi; much better developed plates of the
maxilliped; inner plate of maxilla 1 enlarged and
setose; palps of maxilla 1 and mandible enlarged
These characters suggest a definite generic distinc-
tion we wish not to implement until more material
of Af. zavorus becomes available. Pereopod 5 of Af.
zavorus is depicted strangely, as if it lacks articles
3 and 6 or has a reduced article 6 with tiny dactyl.
Key to the Species of Metaphoxoidrs
(Provisional)
Each lobe of telson with one apical seta, gnathopod I strongly eusirid, palm transverse
? ? ? ? ? ? . M? pftCGfttt
Each lobe of telson with 4 apical setae and spines, gnathopod 1 poorly eusirid, palm weakly
oblique M. xavonu, new species
Metaphoxoidrs picardi Ledoyer
Metaphoxoidrs picardi Ledoyer, 1970:29-31, pi. 8.
DISTRIBUTION.?Madagascar.
Metaphoxoidrs zavorus, new species
Metaphoxus sp.?Griffiths, 1973:296, fig. 10.
DIAGNOSIS.?Metaphoxoides with 4 apical spine-
setae on each lobe of telson; gnathopod 1 poorly
eusirid, palm weakly oblique; palp of mandible
thick, apex of article 3 obliquely excavate, bearing
basofacial setae; inner plate of maxilla 1 large (in
context), bearing 3 setae, palp highly enlarged;
plates of maxilliped well developed.
HOLOTYPE.?Male, 10 mm, as shown in figure 10
of Griffiths, 1973.
TYPE-LOCALITY.?PED 18 V, southwest of Ponta
Zavora, Mozambique, 19 Aug 1964, 25?07'S,
34?34'E, 112 m, dark sandy mud.
REMARKS.?This species is placed provisionally
in this genus; it differs from the type-species, Af.
picardi, in the contrast of characters in the diag-
nosis and eventually should be relegated to a genus
of its own. The species is named for a point of land
near its type-locality.
DISTRIBUTION.?Southern Africa, 112 m.
Diogodias, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
shortened, ventral setae confined apically. Article 1
of antenna 2 not ensiform; article 3 with 2 setules;
facial spines on article 4 in 2 or more rows; article
5 ordinary in size. [PRight mandibular incisor with
3 teeth]; molar absent; pal par hump medium. Palp
of maxilla 1 uniarticulate; inner plate naked.
[PSetation of maxilla 2 weak.] Inner plates of
maxillipeds partly fused together; apex of palp
article 3 not protuberant, dactyl elongate, apical
nail distinct, elongate. Gnathopods dissimilar,
gnathopod 2 moderately to strongly enlarged; arti-
cle 5 of gnathopod 2 either very short, cryptic, or
elongate, of gnathopod 1 elongate and free, with
weak to strong eusirid attachment; palms weakly
oblique or transverse, hands of gnathopods 1-2
weakly elongate or broadened, poorly setose ante-
riorly. Article 5 of pereopods 1-2 setose postero-
proximally. Article 2 of pereopod 3 of broad form,
articles 4-5 of pereopods 3^4 medium to narrow
respectively, article 2 of pereopods 3-4 not setose
posteriorly; pereopod 5 ordinary, article 2 naked
ventrally, article 3 ordinary, dactyl normal. Epi-
mera 1-2 lacking long posterior setae, without
midfacial setae above ventral facial ridge; epimeron
3 of rounded classification and lacking long setae.
Urosomite 1 generally naked; urosomite 3 without
dorsal hook or special process. Peduncle of uropod
1 normally elongate, without apicoventral spike,
without special enlarged apicolateral-medial spine,
[Ppeduncular apices of uropods 1-2 not combed];
inner ramus of uropod 1 naked or with one spine,
no ramus continuously spinose to apex, inner
ramus of uropod 2 ordinary. Uropod 3 ordinary,
elongate article 2 of outer ramus carrying 1-2 short
468 SMITHSONIAN CONTRIBUTIONS IO ZOOLOGY
apical setae. Telson ordinary to elongate, with 1-2
apical spines or setae on each lobe plus setules
[Pwith special lateral setae].
DESCRIPTION.?Rostrum fully developed, elon-
gate. [Fuzz on article 1 of antenna 1 in male un-
known. Calceoli on male antennae 1-2 unknown.
PPrebuccal parts ordinary, poorly separated from
each other, upper lip dominant. PRight lacinia
mobilis unknown;] mandibular palp medium,
article 1 short, article 2 without outer setae, apex
of article 3 oblique. [PLower lip bearing or lacking
cones.] Outer plate of maxilla 1 with 6 spines, no
spine especially thickened. Inner plates of maxil-
liped especially thin, poorly armed. Coxae 2-4 with
weak anterodorsal humps. Some posterior spines on
article 6 of pereopods 1-2 thin and seta-like, others
thick and stiff, midapical spine present. Article 2
of pereopod 5 without facial setae. Peduncle of
iiropod 1 with dorsolateral spines confined apically,
medial spines widely spread; peduncle of uropod 2
with only one medial spine confined apically.
Peduncle of uropod 3 lacking extra subapical setae
or spines. [PTelson with midlateral setules reduced
to one member on each side.]
TYPE-SPECIES.?Metaphoxus longicarpus Ledoyer,
1973 [here selected].
COMPOSITION.?Metaphoxus littoralis Cooper
and Fincham, 1974; Metaphoxus platyrostris
Ledoyer, 1973.
RELATIONSHIP.?The elongate wrist of gnatho-
pod 1 differentiates this genus from Mrtnphoxus.
The rostrum is very elongate. The palpar hump of
the mandible is enlarged in the type and the
flagella of antenna I are elongate. This genus is
named for Diogo Dias, discoverer of Madagascar.
Masculine.
Key to the Species of Diogodias
1. Apex of rostrum spatulate D. ptatyrottris
Apex of rostrum tapering evenly 2
2. Wrist of gnathopod 2 elongate D. littoralis
Wrist of gnathopod 2 short, cryptic D. longitarpus
Diogodias littoralis (Cooper and Fincham),
new combination
Metaphoxus littoralis Cooper and Fincham, 1974:165-168,
figs. 5, 6.
DISTRIBUTION.?New Zealand.
Diogodias longicarpus (Ledoyer), new combination
Metaphoxus longicarpus Ledoyer, 1973:86, pis. 27, 28A.
DISTRIBUTION.?Madagascar.
Diogodias platyrostris (Ledoyer), new combination
Metaphoxus platyrostris Ledoyer, 1973:86, pis. 29, 30.
DISTRIBUTION.?Madagascar.
Vasco, new genus
DIAGNOSIS.?Eyes present but weak. Flagella of
antennae 1-2 unreduced in female. Article 2 of
antenna 1 shortened, ventral setae confined
apically. Article 1 of antenna 2 not ensiforra;
[Particle 3 with 2 setules]; facial spines on article 4
in 2 rows; article 5 ordinary in size. Right mandib-
ular incisor with 3 teeth; molar not triturative,
small, pillow-shaped [possibly lacking spines], not
bearing fuzz; palpar hump medium. Palp of
maxilla 1 uniarticulate; [Pinner plate naked. Seta-
tion of maxilla 2 weak. Inner plate of maxillipeds
ordinary, apex of palp article 3 not protuberant,
dactyl elongate, apical nail distinct, elongate.]
Gnathopods weakly dissimilar, gnathopod 2 mod-
erately enlarged, article 5 of gnathopods 1-2 very
short, cryptic, with eusirid attachment; palms
oblique, hands of gnathopods 1-2 rectangular,
elongate to broadened, poorly setose anteriorly.
Article 5 of pereopods 1-2 scarcely setose postero-
proximally. Article 2 of pereopod 3 of broad form,
articles 4-5 of pereopods 3-4 narrow to medium,
article 2 of pereopods 3-4 not setose posteriorly;
pereopod 5 ordinary, article 2 naked ventrally, arti-
cle 3 ordinary, dactyl normal. [PEpimera 1-2 lack-
ing long posterior setae, without midfacial setae
above ventral facial ridge; epimeron 3 of rounded
NUMBER 245 469
classification and lacking long setae. Urosomite 1
generally naked; urosomite 3 without dorsal hook
or special process.] Peduncle of uropod 1 normally
elongate, without apicoventral spike, without spe-
cial enlarged apicolateral-medial spine, [Ppeduncu-
lar apices of uropods 1-2 not combed]; inner ramus
of uropod 1 naked, no rami continuously spinose to
apex, inner ramus of uropod 2 ordinary. Uropod 3
ordinary, elongate article 2 of outer ramus carrying
2 short apical setae. Telson elongate, with only one
apical spine on each lobe plus setules.
DESCRIPTION.?Rostrum fully developed. [Fuzz
on article 1 of antenna 1 in male unknown.
Calceoli on male antennae 1-2 unknown. PPrebuc-
cal parts ordinary, poorly separated from each
other, upper lip dominant.] Right lacinia mobilis
simple, Habellate; mandibular palp thick, article 1
short (assumed), article 2 without outer setae, apex
of article 3 rounded-transverse. [PLower lip lacking
cones. Outer plate of maxilla 1 with 7 spines, no
spine especially thickened. Inner plates of maxil-
liped especially thin, poorly armed.] Coxae 2-4
without special anterodorsal humps. Some posterior
spines on article 6 of pereopods 1-2 thin and seta-
like, others thick and stiff, midapical seta present,
dactyls stunted. Article 2 of pereopod 5 without
facial setae. Peduncle of uropod 1 with dorsolateral
spines confined apically, medial spines confined
apically; peduncle of uropod 2 with medial spines
or setae widely spread. Peduncle of uropod 3 lack-
ing extra subapical setae or spines. Telson with
ordinary pair of midlateral or dorsal setules on
each side.
TYPE-SPECIES.?Metaphoxus brevidactylus Le-
doyer, 1973 (here selected).
COMPOSITION.?Unique.
REMARKS.?The stunted dactyls of pereopods 1-2
remove this species from Metaphoxus. The diag-
nosis and description do not note that article 5 of
antenna 2 is especially shortened and thin but this
is in relationship to nonmetaphoxin genera be-
cause article 4 is so small in this genus that article
5 appears larger than in other metaphoxin genera.
This genus is named in honor of Vasco da Gama.
Vasco brevidactylus (Ledoyer), new combination
Metaphoxus brevidactylus Ledoyer, 1973:84-86, pi. 26.
DISTRIBUTION.?Madagascar.
Hopiphoxus, new genus
DIAGNOSIS.?Eyes absent. Flagellum of antenna 2
reduced in female. Article 2 of antenna 1 short-
ened, ventral setae confined apically. Article 1 of
antenna 2 not ensiform; article 2 with 2 setules;
facial spines on article 4 in one main row; article 5
short. Right mandibular incisor with 3 teeth;
molar not triturative, medium, pillow-shaped, bear-
ing 2 splayed, semiarticulate spines, bearing fuzz;
palpar hump small. Palp of maxilla 1 uniarticu-
late; inner plate naked. Setation of maxilla 2 weak.
Inner plate of maxillipeds ordinary, apex of palp
article 3 not protuberant, dactyl elongate, apical
nail distinct, short. Gnathopods dissimilar, gnatho-
pod 2 strongly enlarged; article 5 of gnathopods
1-2 very short, cryptic; gnathopods 1-2 with weak
eusirid attachments; palms oblique, hands of
gnathopods 1-2 broadened, poorly setose anteri-
orly. Article 5 of pereopods 1-2 setose postero-
proximally. Article 2 of pereopod 3 of broad form,
articles 4-5 of pereopods 3-4 narrow, article 2 of
pereopods 3-4 not setose posteriorly; pereopod 5
ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Epimera 1-2 lacking long pos-
terior setae, without midfacial setae above ventral
facial ridge; epimeron 3 of rounded classification
and lacking long setae. Urosomite 1 generally
naked; urosomite 3 without dorsal hook or special
process. Peduncle of uropod 1 normally elongate,
without apicoventral spike, without special en-
larged apicolateral-medial spine, [Ppeduncular
apices of uropods 1-2 not combed]; inner ramus of
uropod 1 with one row of marginal setae, outer
ramus shortened, some rami continuously setose to
apex, inner ramus of uropod 2 ordinary. Uropod 3
ordinary, elongate article 2 of outer ramus carrying
2 short to vestigial apical setae. Telson elongate,
with only one apical spine on each lobe.
DESCRIPTION.?Rostrum fully developed. [Fuzz
on article 1 of antenna 1 in male unknown. Cal-
ceoli on male antennae 1-2 unknown. PPrebuccal
parts ordinary, poorly separated from each other,
upper lip dominant. PRight lacinia mobilis bifid,
flabellate;] mandibular palp medium, article 1
short, article 2 without outer setae, apex of article
3 oblique. Lower lip lacking cones. Outer plate of
maxilla 1 with 7 spines, no spine especially thick-
ened. Inner plates of maxilliped especially thin,
poorly armed. Coxae 2-4 without special antero-
470 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
dorsal humps. All posterior spines on article 6 of
pereopods 1-2 thin and seta-like, [Pmidapical spine
or seta present]. Article 2 of pereopod 5 without
facial setae. Peduncle of uropod 1 with dorsolateral
spines confined apically, medial spines weakly
spread; peduncle of uropod 2 with medial setae
weakly spread. Peduncle of uropod 3 lacking extra
subapical setae or spines. Telson with ordinary
pair of midlateral or dorsal setules on each side.
TYPE-SPECIES.?Metaphoxus simillimus J. L.
Barnard, 1967 (here selected).
COMPOSITION.?Unique.
RELATIONSHIP.?Among metaphoxin genera this
genus is unique in the apically setose rami of uro-
pods 1-2, barring the outer ram us of uropod 1.
This genus is named in honor of the Hopi Indians
of North America.
Hopiphoxus simillimus (J. L. Barnard),
new combination
Metaphoxus simillimus J. L. Barnard, 1967:185-138, fig. 66.
DISTRIBUTION.?Baja California, Cedros Trench,
2706 m.
Rikkarus, new genus
DIAGNOSIS OF MALE.?Eyes present. [PFlagella of
antennae 1-2 unreduced in female.] Article 2 of
antenna 1 shortened, ventral setae confined apically.
Article 1 of antenna 2 not ensiform; article 3 with
2 setules; facial spines on article 4 in 2 rows; article
5 thin and short. Right mandibular incisor with
3 teeth; molar not triturative, medium, pillow-
shaped, bearing 1-9 small semiarticulate spines and
enlarged penicillate spine, bearing fuzz; palpar
hump medium. Palp of maxilla 1 uniarticulate;
inner plate naked. Setation of maxilla 2 weak.
Inner plate of maxillipeds ordinary; apex of palp
article 3 not protuberant, dactyl elongate, apical
nail distinct, medium. Gnathopods of medium size;
article 5 of gnathopods 1-2 similar, very short,
cryptic, with eusirid attachment; palms oblique,
hands of gnathopods 1?2 ovatorectangular, elon-
gate, poorly setose anteriorly, article 5 of pereopods
1-2 weakly setose posteroproximally. Article 2 of
pereopod 3 of broad form, articles 4-5 of pereopods
3-4 narrow, article 2 of pereopods 3-4 not setose
posteriorly; pereopod 5 ordinary, article 2 naked
ventrally, article 3 ordinary, dactyl normal.
Epimera 1-2 lacking long posterior setae, without
midfatial setae above ventral facial ridge; epimeron
3 of rounded classification and bearing 3 or fewer
long setae but heavily serrate. L'rosomite 1 bearing
one of more mid ventral bundles of setae; urosomite
3 without dorsal hook or special process. Peduncle
of uropod 1 somewhat shortened, without apico-
ventral spike, without special enlarged apicolateral-
medial spine, peduncular apices of uropods 1-2
combed; inner ramus of uropod 1 with no marginal
spines, no rami continuously spinose to apex,
inner ramus of uropod 2 ordinary. Uropod S ordi-
nary, elongate article 2 of outer ramus carrying
2 short to vestigial apical setae. Telson elongate,
with only one apical spine on each lobe plus
setules, without special dorsal and lateral spines or
setae.
DESCRIPTION.?Rosttuin fully developed; [Fu/z on
article I of antenna 1 in male unknown.] Caheoli
on male primary flagellum of antenna 1 absent,
aesthetascs large and multiple on articles. Calceoli
on article 5 of male antenna 2 present; [Pflagellum
in male without calceoli]. Prebuccal parts ordinary,
poorly separated from each other, upper lip domi-
nant. Right lacinia mobilis bifid, flabellate; man-
dibular palp medium, article 1 short, article 2
without outer setae, apex of article 3 rounded-
transverse. Lower lip bearing cones. Outer plate
of maxilla 1 with 7 spines, no spine especially
thickened. Inner plates of maxilliped especially
thin, poorly armed. Coxae 2-4 without special
anterodorsal humps. All posterior spines on article
6 of pereopods 1-2 thin and seta-like, midapical
spine present. Article 2 of pereopod 5 without facial
setae. Peduncle of uropod 1 with dorsolateral spines
confined apically, medial spines widely spread;
peduncle of uropod 2 with only one medial spine
confined apically, Peduncle of uropod 3 lacking
extra subapical setae or spines. Telson with ordi-
nary pair of middorsal setules on each side highly
medial.
TYPE-SPECIES.?Rikkarus lea, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This genus belongs with the
Metaphoxus group of genera because of the uni-
articulate maxillary palp and poorly setose max-
illa 2. Rikkarus appears dose to Metaphoxus
Bonnier, but differs in the grossly serrate epimeron
3 and in the unexpanded hand of gnathopod 2
NUMBER 245 471
with highly oblique palm, gnathopod 1 duplicating
the size and shape of gnathopod 2 almost precisely.
Rikkarus lea, new species
FICLRES 237-2S9
DESCRIPTION OF MALE (juvenile).?Head about
18 percent of total body length, greatest width
about 45 percent of length; rostrum almost reach-
ing apex of article 3 on antenna 1, unconstricted,
tapering towards apex, apically rounded. Eyes
small, dark, with one layer of clear peripheral
ommatidia. Article 1 on peduncle of antenna 1
about 1.5 times as long as wide, twice as wide as
article 2, apicoventral margin with 7 or more
setules, produced dorsal apex with 3 setules, medial
face lacking fuzz; article 2 about 0.3 times as long
as article 1, apicoventral lateral face with 2 setae;
article 3 slightly overriding article 2 on medial sur-
face; accessory flagellum much shorter than primary
flagellum, 3-articulate; primary flagellum as long
as peduncle, 4-articulate, basal article elongate, all
articles bearing large and numerous aesthetascs.
Spine formula on article 4 of antenna 2 = 3-1, set
2 with one accessory setule, spines thin, dorsomedial
margin fuzzy but elements very elongate and thick,
proximoventral margin with 2-3 penicillate setules,
ventral margin with 4 setae in tandem, ventral
apex with short spine; article 5 about 0.77 times
as long as article 4, facial spine formula = 1, with
accessory setule, dorsodistal corner with 2 spine-
setae, dorsal margin with one calceolus and 2
weakly developed groups of male setae, ventro-
distal apex with one seta and 4 setules, ventral
margin naked; flagellum weakly elongate, about
2.4 times as long as articles 4-5 of peduncle to-
gether, 10-articulate, segments with 1-3 apparent
aesthetascs, no calceoli. Prebuccal complex convex
anteriorly, poorly extended forward, epistome and
upper lip mostly fused together, ventral margin of
upper lip truncate. Palpar hump of mandible
medium to large, broad; right incisor tridentate;
left bidentate with humps; right lacinia mobilis
flabellate, distal branch broad, serrate, face with
cusp, proximal branch simple, longer than distal
branch; left lacinia mobilis broadly flabellate, with
5 teeth; right rakers 2; left 3 plus one rudimentary;
molar formed of weak hump bearing 2 densely
spinulose or penicillate spines; left molar with reg-
ular spination and penicillate spine; palp stout,
article 1 short, article 2 with 3 stout inner distal
setae and one wire-like inner basal seta, article 3
about 0.95 times as long as article 2, rounded-
transverse apex with 9 spine-setae. Each outer lobe
of lower lip with cone, mandibular lobes long,
weakly attenuate, apically rounded, inner lobes al-
most completely fused together (damaged during
dissection). Inner plate of maxilla 1 large, lingui-
form, naked; outer plate with 7 spines; palp reach-
ing apex of outer plate, 1-articulate, bearing 3
apical setae and apicomedial notch. Inner plate of
maxilla 2 shorter and broader than outer plate,
bearing 3 apical setae; subconical outer plate with
one medial, one lateral and 2 apical setae. Inner
plate of maxilliped with one apicomedial spine, one
apicolateral spine and one thinner midapical spine;
outer plate with 3 short medial spines and one
elongate (hidden) basomedial spine; palp article 2
sparsely spinose medially, article 3 unprotuberant,
lacking facial spine, somewhat slipper-shaped and
sparsely spinose medially, dactyl elongate, bearing
medium apical nail and adjacent setule plus second
setule in notch on inner margin. Coxae 1-3 each
with posteroventral cusp and setule near sinus,
coxa 1 expanded distally, with concave anterior
margin, sharp anterior point above anteroventral
bevel (point may be artificial); coxae 2-3 rectangu-
lar; long setal formula of coxae 1-4 = 4-4 4-1;
coxa 4 quadratiform, posterodorsal excavation
medium in size, L-shaped, posterodorsal corner
rounded, ratio of width to length = 17:18. Long
posterior setae on article 2 of gnathopods 1-2 and
pereopods 1-2 = 2-3?4-4, long anteriors = 1 (side)-
0-0-0, short anteriors = 1-2-2-1, no others. Gnatho-
pod 2 scarcely larger than gnathopod 1; article 5
very small and triangular; article 6 elongate, palm
very oblique, defining hump small to medium de-
fining spine large, dactyl fitting palm, posterior
margin of article 6 with one seta; length ratios of
articles 5-6 on gnathopods 1-2 = 31:95 and 32:97,
width ratios = 20:38 and 21:41. Pereopods 1-2
similar to each other; article 4 with one apicofacial
seta, article 5 with 2, largest posterior spine far
exceeding apex of article 6, posterior margin of
article 5 with 4 other similar but shorter spine-
setae laterally, with one setule between spines 3-4,
article 6 naked proximally but with highly distad
set of spine-setae in formula of 2 + 2 plus one mid-
distal spinule, lateral pair of spine-setae short, me-
472 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Cd " ) * *
FIGURE 237.?Rikkanis lea, new species, holotype, juvenile male "a," 2.29 mm.
dial pair as long as dactyl, latter bearing apical
scale but lacking any inner distal setule, notch or
medial pluseta. Coxae 5?7 posteroventral setule for-
mula = 1-1-1. Articles 4-5 of pereopods 3-4 thin,
facial spines absent, formula of facial ridges on
article 2 of pereopods 3-5 = 0-1-1; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 45:18:15:8,
of pereopod 4 = 67:23:16:8, of pereopod 5 =
91:13:12:6, length ratios of pereopod 3 = 61:
22:28:32, of pereopod 4 = 83:50:57=48, of pe-
reopod 5=110:27:21:19; article 2 of pereopod
5 especially large, extending to apex of article 4,
posterior margin with 4 small setule acclivities,
ventral margin with 2; apex of article 6 finely
combed. Epimeron 1 with facial ridge, absent on
epimeron 2, each with rounded subquadrate pos-
teroventral corner, weakly convex posterior mar-
gins with one setule, epimeron 1 otherwise naked,
NUMBER MB
G2t
FIGURE 238.?Rikkarus lea, new species, holotype, juvenile male "a," 229 mm.
epimeron 2 with 3 facial setae in tandem horizon-
tally, epimeron 3 with 6 teeth on lower half of pos-
terior margin, upper tooth blunt, others sharp,
becoming small and cusp-like ventrally, each with
setule or seta in adjacent sinus, epimeron 3 other-
wise naked. Urosomites naked, unproduced dor-
sally. Peduncle of uropod 1 naked basofacially,
lateral apex, medial apex and medial margin with
one spine, rami of uropods 1-2 naked dorsally,
apices of inner ramus on uropod 1 and both rami
of uropod 2 with sharp, attenuate scale and barest
indication of completely submerged and fused nail,
outer ramus of uropod 1 slightly shortened and
thinner than inner ramus, curved, apex simple,
blunt, inner rami of uropods 1-2 lacking baso-
medial setule, peduncle of uropod 2 with 3 dorsal
spines, medially with one apical spine, lateral pe-
duncular apices of uropods 1-2 very minutely and
474 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
FIGURE 239.?Rikkarus lea, new species, holutype, juvenile male "a." 2.29 mm.
weakly combed. Uropod 3 scarcely elongate, pe-
duncle with 3 ventral spines, one dorsolateral spine,
one dorsomedial setule; rami submasculine, inner
extending to M. 87 on article 1 of outer ramus,
with 2 long apical and 2 medial setae, article 2
of outer ramus elongate, 0.57, bearing 2 apical set-
ules, lateral margin of article 1 naked except for
2 apicolateral spines, medial margin with one long
seta, medial apex with one long seta, one short
spine. Telson elongate, not fully cleft, length-
width ratio = 18:13, apices tapering, of medium
breadth to narrow, subtruncate, bearing one long_
spine and one apicolateral setule, dorsal pair of
plusetules alike, near middle of lobe, elongate.
Small bulbar setules of cuticle extremely sparse,
emergent setules very short, scarcely as long as
width of pore.
ILLUSTRATIONS.?Lateral view of head drawn from
slightly dorsal aspect, dorsal view of head recon-
structed; unique juvenile male with small penial
processes.
HOLOTYPE.?AM, immature male "a," 2.29 mm.
Unique.
TYPE-LOCALITY.?SBS 3, 23 Jul 1973, off Mala-
bar, New South Wales, Australia, 150? 16' E,
33?58'42" S, 66 m.
MATERIAL.?AM, one sample (I).
DISTRIBUTION.?New South Wales, off Malabar,
66 m.
Jopara, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 reduced in female. Article 2 of antenna 1 ordi-
nary, ventral setae confined apically. Article 1 of
antenna 2 not ensiform; article 3 with 2 setules;
facial spines on article 4 in 2 rows; article 5 espe-
cially short. Right mandibular incisor with 3 teeth;
molar not triturative, medium, elongate, conical,
bearing 4 or more splayed, semiarticulate spines,
bearing fuzz; palpar hump small. Palp of maxilla 1
uniarticulate; inner plate naked. Setation of
maxilla 2 weak. Inner plate of maxillipeds partly
fused together, apex of palp article 3 not protuber-
ant, dactyl elongate, apical nail distinct, medium.
Gnathopods dissimilar, gnathopod 2 strongly en-
larged; article 5 of gnathopod 2 very short, cryptic,
elongate on gnathopod 1, with eusirid attachment,
palms oblique to transverse to chelate, hands of
gnathopods 1-2 rectangular, broadened, poorly
setose anteriorly. Article 5 of pereopods 1-2 setose
posteroproximally. Article 2 of pereopod 3 of
NUMBER 245 475
broad form, articles 4-5 of pereopods 3-4 narrow
to medium, article 2 of pereopods 3-4 not setose
posteriorly; pereopod 5 ordinary, article 2 naked
ventrally, article 3 ordinary, dactyl normal. Epi-
mera 1-2 lacking long posterior setae, without mid-
facial setae above ventral facial ridge; epimeron 3
of rounded classification and lacking long setae.
Urosomite 1 generally naked; urosomite 3 without
dorsal hook or special process. Peduncle of uropod
1 normally elongate, without apicoventral spike,
without special enlarged apicolateral-medial spine,
peduncular apices of uropods 1-2 combed, inner
ram us of uropod 1 with marginal spines in one
row, no rami continuously spinose to apex, inner
ramus of uropod 2 ordinary. Uropod 3 small to
minute, lacking article 2 of outer ramus, rami
minute, shorter than peduncle, extending equally,
in female each carrying 1-2 vestigial apical setae.
Telson ordinary, with 1-2 apical spines or setae on
each lobe plus setules.
DESCRIPTION.?Rostrum small to obsolescent.
Fuzz on article 1 of antenna 1 in male present;
calceoli on male primary flagellum of antenna 1
present. Calceoli on article 5 of male antenna 2
present; flagellum in male with calceoli. Prebuccal
parts strongly extended forward, massive, distinct,
both epistome and upper lip prominent. Right
lacinia mobilis bifid, flabellate; mandibular palp
thin, article 1 elongate, article 2 without outer
setae, apex of article 3 oblique. Lower lip lacking
cones. Outer plate of maxilla 1 with 7 spines, no
spine especially thickened, one attached medially
and pointing medially. Inner plates of maxilliped
poorly armed, thick. Article 3 of gnathopod 1 elon-
gate. Coxae 2-4 without special anterodorsal
humps. Some posterior spines on article 6 of pereo-
pods 1-2 thin and seta-like, others thick and stiff,
midapical spine or seta absent. Article 2 of pereo-
pod 5 without facial setae. Peduncle of uropod 1
with dorsolateral spines confined apically, medial
spines widely spread; peduncle of uropod 2 with
only one medial spine confined apically. Peduncle
of uropod 3 lacking extra subapical setae or spines.
Telson with pair of midlateral or dorsal setules on
each side, highly apical.
TYPE-SPECIES.?Japara papporus, new species
(here designated).
COMPOSITION.?Unique.
RELATIONSHIP.?The type-species of this genus
has general resemblance to Metaphoxus fultoni
(Scott) because of the shape of gnathopod 1; the
genus appears close to Metaphoxus because of the
uniarticulate palp of maxilla 1, the nontriturative
molar, broad article 2 of pereopod 3, unproduced
article 3 of the maxillipedal palp and the general
appearance of the head, body, coxae, and pereo-
pods. Japara differs from Metaphoxus in the minia-
turized rami of uropod 3 and the short, broad
inner plates of the maxillipeds. In terms of general
knowledge about Phoxocephalidae we suggest that
the condition of the maxillipeds represents a prim-
itive state and the condition of the third uropods
an advanced state in relation to Metaphoxus. The
second maxilla of Metaphoxus, Leptophoxus Sars,
and Japara also shares similarities in the numerical
reduction of setae.
Japara differs from Kondoleus in the absence of
spouts on the maxillipeds, the more elongate dac-
tyl of the maxillipeds, the normally subchelate
gnathopod 2, the thin pereopod 4, the much shorter
rami of uropod 3 and the absence of long distal
setae on uropods 1-2.
Japara papporus, new species
FIGURES 240-242
DESCRIPTION OF FEMALE.?Head about 16 per-
cent of total body length, greatest width about 75
percent of length; rostrum obsolete, broad, scarcely
reaching along article 1 on antenna 1. Eyes small,
largely occluded with pigment, ommatidia espe-
cially large. Article 1 on peduncle of antenna 1
about 1.33 times as long as wide, about 1.5 times
as wide as article 2, ventral margin with 2 setules,
ventral apex with 2 long setules, strongly produced
dorsal apex with 2 setules; article 2 about 0.75
times as long as article 1, with apicoventral cycle
of 4 setae; primary flagellum with 4 articles, about
as long as article 2 of peduncle, bearing large aes-
thetascs; accessory flagellum with 3 articles. Spine
formula on article 4 of antenna 2 = 1-2-1, dorsal
margin naked, ventral margin with 3 ventrodistal
medium to short spines and 4 basofacial setules;
article 5 about 0.60 times as long as article 4, facial
spine formula = 1, dorsal apex bearing 2 large
spines, ventral margin with 2 sets of setules, 4 ven-
trodistal short to medium spines and setae; flagel-
lum not half as long as articles 4-5 of peduncle
combined, with 4-6 articles. Mandibles with weak
476 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 240.?Japara papporus, new species, holotype, female "a," 2.52 mm (y ? male "y,1
2.64 mm).
NUMBER 245 477
palpar hump; right incisor with 3 teeth; left incisor
with 5 humps in 2 branches; right lacinia mobilis
broadly flabellate, subbifid, denticulate, with one
facial hump; left lacinia mobilis with 6 teeth; right
rakers 2; left rakers 4; molar in form of bulbous
cone, with minute cusps, large spine, setules, plu-
setule and fuzz; palp article 1 elongate, article 2
with one short inner seta in middle, article 3 as
long as article 2, oblique apex with 3 spine-setae,
basofacial formula = 0. Inner plate of maxilla 1
ordinary, conical, naked, palp uniarticulate, with
2 apical and one lateral setae. Plates of maxilla 2
extending subequally, of subequal width, setae re-
duced, outer with one apicolateral seta, inner with
2 medial setae. Inner plate of maxilliped with one
large, thick apical spine, 2 apicofacial setae, no
medial setae; outer plate with 4 medial spines, no
apicolateral setae; palp articles 1-3 lacking apico-
lateral setae, medial margin of article 2 weakly se-
tose, article 3 unprotuberant, with one facial seta,
nail of article 4 medium, with 2 accessory setules.
Coxa 1 scarcely expanded distally, anterior margin
weakly convex; main ventral setae of coxae 1-4 =
0-0-0-0; anterior and posterior margins of coxa 4
almost parallel, posterior margin convex, postero-
dorsal corner sharp-rounded, posterodorsal margin
short, concave, width-length ratio of coxa 4 = 6:7.
Long posterior setae on article 2 of gnathopods 1-2
and pereopods 1-2 = (l-2)-2-2-4, short anteriors
not counted, short posteriors = (l-2)-2-2-2. Gna-
thopods enlarged; article 3 of gnathopod 1 elongate,
wrist-hand attachment eusirid, hand rectangular,
article 3 of gnathopod 2 short, hand almost rectan-
gular, slightly larger than hand of gnathopod 1;
with ratios of articles 3-6 on gnathopods 1-2 =
14:16:25:47 and 23:22:20:61, length ratios =
31:30:62:80 and 18:35:28:86; palmar humps large,
palms 1 and 2 parachelate and oblique respec-
tively; article 5 of gnathopod 1 elongate, poste-
rior margin lobate, article 5 of gnathopod 2 short,
triangular, lobate, cryptic. Pereopods 1-2 similar;
weakly setose, facial setae formula on article 4 =
0 and 0, on article 5 = 0 and 0, main spine of arti-
cle 5 extending to M. 70 on article 6, article 5
lacking proximoposterior spines, spine formula of
article 6 = 2 and 1, spines especially short; acclivity
on inner margin of dactyls of pereopods 1-2 ab-
sent, marked by slit, lacking emergent setule, mid-
facial pluseta ordinary, highly proximal and dor-
sal. Coxae 5-7 posteroventral setule formula =
1-1-1. Articles 4-5 of pereopod 3 of ordinary width,
of pereopod 4 narrow, facial spine rows absent,
facial ridge formula on article 2 of pereopods
3-5 = 0-1-1, pereopods 3-5 short; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 36:28:25:11,
of pereopod 4 = 60:20:15:9, of pereopod 5 =
90:20:19:9, length ratios of pereopod 3 = 55:
26:32:23, of pereopod 4 = 80:28:40:25, of pe-
reopod 5=112:20:22:20; article 2 of pereopod
5 reaching apex of article 4; medial apex of
article 6 finely combed, lacking digital processes.
Posteroventral corners of epimera 1-2 rounded,
posterior margins convex, posteroventral setae ab-
sent, corner of epimeron 3 weakly protuberant,
with setule sinus, posterior margin straight, with
one setule notch, ventral margin naked, epimera
1-3 with setule on posterodorsal margin set in
weak notch, epimeron 3 with total of 3 posterior
setules. Urosomite 1 tall, naked, articulation line
complete; urosomite 3 weakly protuberant dorsally.
Rami of uropods 1-2 with articulate, enlarged api-
cal nails, inner ramus of uropod 1 with one dorsal
spine, other rami naked; peduncle of uropod 1
with one apicolateral spine and no basofacial setae,
medially with 2 marginal spines, apicalmost en-
larged, peduncle of uropod 2 with one dorsal spine,
medially with one thin apical spine, apicolateral
corners of peduncles on uropods 1-2 with weak
comb. Uropod 3 reduced, peduncle with one ven-
tral spine, dorsally with lateral spinule, one medial
spine; rami scarcely half as long as peduncle, coni-
cal, extending equally, each apex with 1-2 setules,
otherwise naked, article 2 of outer ramus absent.
Telson ordinary, length-width ratio =1:1, not
fully cleft, each apex narrow, truncate, lateral ac-
clivity absent, bearing 2 short setules, each subapex
with 2 long, diverse plusetules, possibly represent-
ing ordinary midlateral setules. Cuticle with widely
scattered pipes throughout, surface bearing fine
striations in form of linear fingerprint pattern.
OBSERVATIONS (female).?Article 2 of antenna 1
overriding article 3 medially, article 3 of normal
shortness. Article 2 of antenna 2 largely fused to
neighboring articles. Inner plate of maxilla 1 com-
posed of folded halves forming a cone. Inner plates
of maxillipeds apically truncate but nodulose.
Short posterior setae of article 2 on gnathopods
1-2 and pereopods 1-2 straw-like. Lobes of telson
highly arched and forming ventral channel for
apposition of uropod 3.
478 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 241.?Japara papporus, new species, holotype, female "a," 2-52 mm (y ? male "y," 2.64
mm; K ?? spine of palm on gnathopod 2).
NUMBER 245 479
FIGURE 242.?Japara papporus, new species, holotype, female "a," 2-52 mm (m ? male "m,
2-86 mm; y ? male "y," 2-64 mm; Y = denticles; pereopod 3 enlarged, see text).
480 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Eggs: Holotype, female "a," 2.52 mm, with 2
large eggs; female "e," 2.61 mm, 2 large eggs; fe-
male "f," 2.32 mm, one large egg.
DESCRIPTION OF MALE.?Following parts like
those of female: prebuccal, mandibles (except
palps), lower lip, maxillae, maxilliped, gnathopod
1, gnathopod 2 (but hand slightly stouter than in
female), pereopods 1-4, uropods 1-2. Parts unlike
those of female: eyes larger than in female, omma-
tidia free of occluding pigment, brass in color.
Article 1 of antenna 1 with medial patch of fuzz;
primary flagellum with 5 articles, first 2 articles
each with large calceolus; accessory flagellum 3-
articulate. Antenna 2 basally stouter than in fe-
male, anterior margins of articles 3-4 with long
fuzz, anterior margin of article 5 with male setule
tufts and 0-2 calceoli, facial spine formula of arti-
cle 4 = 2-2, of article 5 = 1; flagellum highly elon-
gate, about 19-articulate, flagellar formula = 19, 1,
2, 4, 6 . . . 18. Article 3 of mandibular palp about
1.2 times as long as article 2, bearing 4 apical spine-
setae and with 2 basofacial setae, one on each side.
Long posterior setae of article 2 on gnathopods 1-2
and pereopods 1-2 = 1-1-1-2. Coxae 1-4 all
slightly narrower than in female. Pereopod 5 with
narrower and slightly less extended article 5, arti-
cles 4-5 narrower and article 5 shorter than in
female, article 5 with sloping anterodistal margin
bearing 3 clavate and curved spines and 2 long
undulant setae, anterior margin of article 6 bearing
one long seta. Epimeron 1 with broadly rounded
posterior margin, epimera 2-3 longer anteroposte-
riorly than in female, epimeron 2 with one facial
seta, epimeron 3 with 4 posterior setule notches.
Urosomite 1 with one setule on each side at base
of uropod 1. Rami of uropod 3 larger than in fe-
male but only about two-thirds as long as peduncle,
broader, more strap-shaped and less conical, bear-
ing long setae. Telson with row of dorsal denticles
on each lobe, most of these denticles reminiscent
of rose thorns but those at ends of row more chisel-
shaped, apex of each lobe with additional long
spine seta besides 2 short setules found in female.
Cuticular pipes much sparser than in female but
bulbar setules, though sparse, more numerous than
in female.
ILLUSTRATIONS.?View of dissected female pereo-
pod 3 drawn about 1.4 times larger than views of
female pereopods 4 and 5; one spine on outer plate
of maxilliped hidden.
HOLOTYPE.?NMV, female "a," 2.52 m. (Carcass
of holotype, comprising pereonites 1-7 minus ap-
pendages, lost during transferral to repository.)
TYPE-LOCALITY.?CPBS 41N/1171. 1 Nov 1971,
Western Port, Victoria, Australia, 12.8 m, silty
sand, gravel.
VOUCHER MATERIAL.?Type-locality: "female "e,"
2.61 mm; female "(," 2.32 mm; CPBS 33N/5: male
"y," 2.64 mm (illus). CPBS 23N/2: male "m," 2.86
mm (illus.); female "d," 2.88 mm. SBS Long Reef:
male "g," 2.42 mm; female "h," 2.28 mm; juvenile
"')," 1.37 mm.
MATERIAL.?CPBS, 39 samples from 9 stations
(23); RHM, one sample (1); SBS, one sample (1).
DISTRIBUTION.?Victoria, Western Port, neritic
and 10-13 m, gravel, sand or shell. New South
Wales, Long Reef, N of North Head, Sydney,
43 m.
Kondoleus, new genus
DIAGNOSIS.?Eyes present. Flagella of antenna 1-2
reduced in female. Article 2 of antenna 1 ordinary,
ventral setae confined apically. Article 1 of antenna
2 not ensiform; article 3 with 2 setules; facial
spines on article 4 circular and spherical, in 2
rows; article 5 especially short. Right mandibular
incisor with 5 teeth; molar not triturative, medium,
conical, bearing 3 long clumped spines with partly
common base, one spine bearing fuzz; palpar hump
small. Palp of maxilla 1 uniarticulate; inner plate
naked. Setation of maxilla 2 weak. Inner plates of
maxillipeds short, bearing basal spouts pointing
laterally; apex of palp article 3 not protuberant,
dactyl short, stubby, apical nail distinct, mostly
immersed, medium. Gnathopods dissimilar, gnatho-
pod 2 moderately enlarged; article 5 of gnathopod
2 short, cryptic, of gnathopod 1 elongate, with
eusirid attachment, palms chelate, hands of gnath-
opods 1-2 broadened, poorly setose anteriorly. Arti-
cle 5 of pereopods 1-2 setose (or spinose) postero-
proximally. Article 2 of pereopod 3 of broad form,
articles 4-5 of pereopods 3-4 broad, oddly trape-
zoidal, article 2 of pereopods 3-4 not setose poste-
riorly; pereopod 5 ordinary, article 2 naked ven-
trally, article 3 ordinary, dactyl normal. Epimera
1-2 lacking long posterior setae, without midfacial
setae above ventral facial ridge; epimeron 3 of
rounded classification and lacking long setae. Uro-
somites 1-2 with large dorsal and glandular humps,
NUMBER 245 481
generally naked except for sparse apicoventral setae
near base of uropod 1; urosomite 3 without dorsal
hook or special process. Peduncle of uropod 1 nor-
mally elongate, without apicoventral spike, without
special enlarged apicolateral or medial spine, uro-
pods 1-2 with occasional seta, peduncular apices
of uropods 1-2 not combed, inner ramus of uropod
1 with marginal spines in one row, one ramus con-
tinuously setose to apex, inner ramus of uropod 2
especially shortened, partially fused to peduncle.
Uropod 3 small, article 2 of outer ramus carrying
2 long apical setae. Telson elongate, with 2 apical
spines or setae on each lobe plus setules.
DESCRIPTION.?Rostrum obsolescent. Fuzz on
article 1 of antenna 1 in male present; calceoli on
male primary flagellum of antenna 1 absent. Cal-
ceoli on article 5 of male antenna 2 absent; flagellum
in male with calceoli. Prebuccal parts ordinary,
massive, strongly distinct, upper lip domi-
nant. Right lacinia mobilis bifid, flabellate; man-
dibular palp thin, article 1 elongate, article 2 with-
out outer setae, apex of article 3 oblique. Lower
lip bearing cones. Outer plate of maxilla 1 with
7 spines, no spine especially thickened. Inner plates
of maxilliped especially thin, poorly armed. Coxae
2-3 tapering distally, coxae 2-4 without special
anterodorsal humps. All posterior spines on article
6 of pereopods 1-2 thick and stiff, midapical spine
present. Article 2 of pereopod 5 without facial
setae. Peduncle of uropod 1 with dorsolateral
spines confined apically, medial spines widely
spread; peduncle of uropod 2 with only one medial
setule confined apically; peduncle of uropod 3
lacking extra subapical setae or spines. Telson with
pair of midlateral or dorsal setules on each side.
TYPE-SPECIES.?Kondoleus tekin, new species
(here selected).
COMPOSITION.?Unique.
RELATIONSHIP.?Despite similarities to Joubinella
Chevreux in gnathopods and flagellum of antenna
2, Kondoleus appears more closely approximate to
the Metaphoxus group of genera by virtue of man-
dibular molar and first maxillary palp. Kondoleus
differs from both Metaphoxus and Joubinella in
the stubby article 4 of the maxillipedal palp, the
shortened inner rami of uropod 2 and further
differs in numerous minor characters herein de-
scribed for the type-species of Kondoleus and not
heretofore attributed to generic level, to wit: a
huge glandular mass in urosomites 1-2, with dorsal
ducts on those segments; salivary spouts at the base
of the inner plates on the maxillipeds; and oddly
shaped triangular or trapezoidal articles 4-5 of
pereopods 3-4.
This genus is distinguished from Japara, new
genus, in the large rami of uropod 3, and the
stubby palp article 4 of the maxilliped among nu-
merous characters ranked secondarily (see illustra-
tions and descriptions).
Kondoleus bears considerable resemblance to
Limnoporeia and should be studied in detail for
its possible origins near that genus. Kondoleus
differs from Limnoporeia in its heavily modified
antennae, maxillipedal spouts, and stubby maxilli-
pedal dactyl, in the heavy reduction of the rostrum,
presence of setae on uropods 1-2, shortened inner
ramus of uropod 2, and a host of other specializa-
tions.
Kondoleus tekin, new species
FIGURES 243-246
DESCRIPTION OF FEMALE.?Head about 15-17 per-
cent of total body length, greatest width about 75
percent of length; rostrum obsolescent, broad, not
reaching along article 1 on antenna 1. Eyes me-
dium, largely occluded with pigment, ommatidia
especially large. Article 1 on peduncle of antenna
1 about 1.3 times as long as wide, about 1.6 times
as wide as article 2, apicoventral margin with 3
setules, unproduced dorsal apex with 2 setules;
article 2 0.9-1.1 times as long as article 1, apico-
ventral corner produced, with 2 long setules; article
3 enlarged; primary flagellum with 3 articles, about
0.75 times as long as peduncular article 3, bearing
large aesthetascs; accessory flagellum with 2 articles.
Antenna 2 short and thick; article 3 with one facial
seta; spine formula of article 4 = 7^4 or 9-4, some-
times distal group appearing divided into subequal
groups, these spines nearly spheroidal, very short
and almost fully dorsal, ventroproximal face with
3 setules, ventral margin naked, ventrodistal corner
with one seta; article 5 about 0.45 times as long as
article 4 and narrower, spine formula = 2-1 or
simply 3, these spines spheroidal and dorsal, ventral
margin with one acclivity and seta, ventrodistal
margin with one stout spine and seta; flagellum
2-articulate, very short, basal article almost fully
immersed in peduncle. Mandibles with weak-
482 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 243.?Kondoleus tekin, new species, holotype, female "a," 3.00 mm (m ? male "m,"
3.02 mm).
NUMBER 245 483
FIGURE 244.?Kondoleus tekin, new species, holotype, female "a," 3.00 mm (m = male "m,
3.02 mm; n ? female "n," 3.06 mm).
medium pal par hump; right incisor with 6 teeth
and humps, left incisor with 6 humps in 2
branches; right lacinia mobilis broadly flabellate,
subbifid, marked by sigmoid notch, denticulate,
with 1-2 facial humps; left lacinia mobilis with 6
teeth; right rakers 2; left rakers 3 plus one rudi-
mentary; molar in form of elongate protrusion de-
marcated mainly by spines, each molar with 3 pri-
marily long spines, one of these plumed; palp article
1 elongate, article 2 naked, article 3 about as
long as article 2, oblique apex with 4 spine-setae,
midfacial formula = 0-1. Inner plate of maxilla 1
long, narrow, naked; palp uniarticulate, short, with
3 apicalmedial and lateral setae. Inner plate of
maxilla 2 shorter than outer, outer scarcely broader
than inner, outer with one apicolateral seta, inner
with one medial seta, plates short and poorly se-
tose. Inner plate maxilliped with 2 large, thin
apical spines, otherwise naked, bases produced into
laterally directed spouts; outer plate with 5 medial
484 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCI
FIGURE 245.?Kondoleus tekin, new species, holotype, female "a," 3.00 mm (m ? male "m,1
3.02 mm).
spines (2 forming pair); palp articles 1-2 naked
apicolaterally, medial margin of article 2 weakly
setose, article 3 with 3 facial setae in triangle, no
lateral setae, article 4 short, stubby, nail medium,
with 2 accessory setae. Coxa 1 expanded apico-
posteriorly, anterior margin convex, coxae 2-3
weakly pointed; main ventral setae of coxae 1?4 =
0, setules = 3-3-3-2; anterior and posterior mar-
gins of coxa 4 strongly divergent, ventral margin
short, rounded, posterior margin very oblique, al-
NUMBER 245 485
most straight, posterodorsal corner rounded, pos-
terodorsal margin short, concave, width-length
ratio of coxa 4 = 13:14. Long posterior setae on
article 2 of gnathopods 1-2 and pereopods 1-2 =
0-(3-4)-6-(5-6), short anteriors = 0-2-1-1, long
anteriors = 0-2-4-(2-3), no others. Gnathopods en-
larged, article 6 broad, parachelate, hand-wrist at-
tachment eusirid, gnathopod 2 stouter than gnatho-
pod 1; width ratios of articles 5-6 on gnathopods
1-2 = 22:52 and 16:60, length ratios = 72:82
and 30:95; palmar humps ordinary; article 5 of
gnathopod 1 elongate, posterior margin rounded,
almost lobate, article 5 of gnathopod 2 short, cryp-
tic. Pereopod 1 longer than pereopod 2, especially
article 4, posterior margin of latter poorly setose,
anterodistal setae formula on article 4 = 4 and 3,
on article 5 = 1 and 1, main spine of article 5 ex-
tending to M. 20 on article 6, article 5 with pair
of proximoposterior spines and poorly setose, spine
formula of article 6 = 3 + 1 plus middistal spine,
some spines especially short, acclivity on inner mar-
gin of dactyls of pereopods 1-2 obsolescent, emer-
gent setule long, midfacial pluseta highly proximal.
Coxae 5-7 posteroventral setule formula = 1-1-1.
Articles 4-5 of pereopods 3-5 broad, trapezoidal,
facial spine rows absent, facial ridge formula on
article 2 of pereopods 3-5 = 0-0-0; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 68.61:60:9,
of pereopod 4 = 71:53:41:10, of pereopod 5 =
108:35:29:10, length ratios of pereopod 3 =
72:30:39:22, of pereopod 4 = 92:32:38:22, of pe-
reopod 5 = 128:60:41:18; article 2 of pereopod
5 exceeding apex of article 7 because articles
4-7 flexed backward permanently, article 2 huge,
suborbicular, smooth; article 4 of pereopod 5
strongly extended along article 5 posteriorly, also
true for extension of article 5 along article 6 on
pereopods 3-4; article 6 very short on pereopods
3-5, medial apex of article 6 smooth. Posteroven-
tral corner of epimera 1-2 rounded, posterior mar-
gin straight to convex, posteroventral corner of
epimeron 3 rounded-quadrate, posterior margin
straight, epimera 1-3 naked except for setule on
posterodorsal margin. Urosomite 1 with lateral set-
ule at base of uropod 1, articulation line incom-
plete, short, dorsal urosomite 1 hugely elevated and
posterodorsally extended as hump bearing ducts
from large gland filling urosomites 1-2 and part
of pleonite 3, urosomite 2 also with hump, uroso-
mite 3 weakly protuberant dorsally. Rami of uro-
pods 1-2 with articulate apical nails, outer ramus
of uropod 1 with one dorsal spinule, inner with 2
dorsal and one ventral elongate setae, outer ramus
of uropod 2 with one long dorsal seta, inner ramus
short, essentially fused to peduncle, with one me-
dium dorsal seta; peduncle of uropod 1 with one
apicolateral spinule and no basofacial setae, medi-
ally with 2 marginal spines, apicalmost an ordinary
spine, peduncle of uropod 2 with one long dorso-
distal seta, medially with one small apical seta,
apicolateral corners of peduncles on uropods 1-2
lacking comb. Peduncle of uropod 3 with one ven-
tral setule, dorsally with one medial setule, later-
ally naked; rami feminine, inner extending to M.
45 on article 1 of outer ramus, apex with one seta,
medial and lateral margins naked, article 2 of outer
ramus short, 0.08, bearing 2 long setae, apicomedial
margin of article 1 with 2 setae, lateral margin
with 2 pairs of setae. Telson especially long, length-
width ratio = 14:11, not fully cleft, each apex nar-
row, rounded, lateral acclivity absent, bearing short
lateral setule, seta next medial elongate, next seta
medium, midlateral setules large, diverse. Cuticle
covered with dense polygonal structure formed by
concentrations of villi and pebbles, bulbar setules
rarely present, polygons especially heavy on coxae,
article 2 of pereopods 3-5, midbody, antenna 2,
weaker on epimera and antenna 1, in other places
villi not organized into polygonal structure, dimin-
ished or absent on urosome, head, apices of gnatho-
pods and pereopods.
DESCRIPTION OF MALE.?Following parts like
those of female: head shape, epistome, upper lip,
right molar, lower lip, maxillae, maxilliped, gnatho-
pods, pereopods 1-2, epimera, medial aspect of
uropod 1, length of telson. Eyes larger than in fe-
male, clear of occluding pigment, gold in color in
preservative. Head about 15 percent of total body
length. Antenna 1 with brush of thick setules on
lateral (unusual) surface of article 1, aesthetascs
more densely developed than in female; primary
flagellum 4-articulate. Antenna 2 scarcely thicker
than in female; flagellum elongate, flagellar for-
mula = 21, 1-4, 6, 8 , . . . 20 or 15, 1, 3, 4, 6, 8 . . . 12.
Spine formula on article 4 of peduncle either 7-4
or 7-5, article 5 with either 3 or 4 dorsal spines.
Right lacinia mobilis with sigmoid cusp and notch
seen in female straightened to right angles; left
lacinia mobilis with subdenticulations; right and
left rakers = 3 and 4; left molar widi 2 spines (one
486 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCf
FIGURE 246.?Kondoleus tekin, new species, holotype, female "a," 3.00 mm (m ? male "m,
3.02 mm; K - denticle).
NUMBER 245 487
ciliate); mandibular palp article 3 with 2 basofacial
setae on same face, apex with 4 setae, inner apical
acclivity and one subapical seta; slight differences
from female in coxa 1 (see illustration). Pereopods
3-5 with weakly distinctive shapes on several arti-
cles (see illustrations). Long posterior setae on arti-
cle 2 of gnathopods 1-2 and pereopods 1-2 = 0 -3 -
(4-5)-5, long anterior setae = 0-3-5-3. Coxa 4 rela-
tively slightly smaller in relation to coxa 1 than
is coxa 4 to coxa 1 in female, similar statement for
male coxae 2-3. Dorsal protuberances of urosomites
1-2 somewhat more sharply defined than in female.
Uropod 1 with one long basofacial seta; outer
ram us of uropod 2 with one or pair of setae dor-
sally, inner ramus very broad and scaliform. Inner
ramus of uropod 3 reaching to M. 67 on outer
ramus, both rami with numerous long setae except
on outer margin of inner ramus. Telson with row
of medial denticles on both lobes, apex of each
lobe with one long seta in middle, with lateral
short bulbar seta and short medial seta.
ILLUSTRATIONS.?Left mandibular molar of fe-
male with 2 spines broken off as shown by 4 basal
lines depicting nerve ducts.
HOLOTYPE.?NMV, female "a," 3.00 mm.
TYPE-LOCALITY.?CPBS B3/5a, 29Sep 1964, West-
ern Port, Victoria, Australia, 11 m, sand.
VOUCHER MATERIAL.?Type-locality, female "n,"
3.06 mm (illus.); CPBS 25S/5, male "m," 3.02 mm
(illus.); CPBS A3/5, male "e," 2.91 mm.
MATERIAL.?CPBS, 8 samples from 6 stations
(10).
DISTRIBUTION.?Victoria, Western Port, 8.2-18.3
m, sand, sandy mud.
Limnoporeia Fearn-Wannan
Limnoporeia Fearn-Wannan, 1968:37-38.
DIAGNOSIS.?Eyes present. Flagellum of antenna
2 reduced in female. Article 2 of antenna 1 short-
ened to ordinary to elongate, ventral setae confined
apically. Article 1 of antenna 2 not ensiform; arti-
cle 3 with 2 setules; facial spines on article 4 in 2
or more rows, plus special apical spines; article 5
thin and short. Right mandibular incisor with 3
teeth; molar not triturative, medium, elongate,
conical, bearing two or fewer conical fused spines,
often with button, bearing fuzz; palpar hump small
to large. Palp of maxilla 1 uniarticulate; inner
plate naked. Setation of maxilla 2 weak. Inner
plate of maxillipeds ordinary; apex of palp article
3 not protuberant, dactyl elongate, apical nail dis-
tinct, elongate. Gnathopods large, dissimilar, gna-
thopod 2 strongly enlarged; article 5 of gnathopod 1
elongate to ordinary, free, of gnathopod 2 almost
cryptic, with weak eusirid attachment; palms che-
late, hands of gnathopods 1-2 elongate, poorly se-
tose anteriorly. Article 5 of pereopods 1-2 setose
posteroproximally. Article 2 of pereopod 3 of broad
form, articles 4-5 of pereopods 5-A narrow, article
2 of pereopods 3-4 not setose posteriorly; pereopod
5 ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Epimera 1-2 lacking long pos-
terior setae, without midfacial setae above ventral
facial ridge; epimeron 3 of rounded classification
and bearing 3 or fewer long setae. Urosomite 1
generally naked; urosomite 3 without dorsal hook
or special process. Peduncle of uropod 1 normally
elongate, without apicoventral spike, without spe-
cial enlarged apicolateral or medial spine, pedun-
cular apices of uropods 1-2 combed; inner ramus
of uropod 1 with marginal spines in one row, no
rami continuously spinose to apex, apical nails ab-
sent, inner ramus of uropod 2 ordinary. Uropod 3
ordinary, elongate article 2 of outer ramus carry-
ing 2 short to long apical setae. Telson elongate,
with 1-4 apical spines or setae on each lobe plus
setules, without special dorsal and lateral spines
or setae.
DESCRIPTION.?Rostrum fully developed, often
with apicoventral process. Fuzz on article 1 of an-
tenna 1 in male present; calceoli on male primary
flagellum of antenna 1 present. Calceoli on article
5 of male antenna 2 present; flagellum in male
with calceoli. Prebuccal parts strongly extended
forward, massive, strongly distinct, upper lip domi-
nant. Right lacinia mobilis bifid, flabellate; man-
dibular palp of medium size, article 1 short,
article 2 without outer setae, apex of article 3
oblique. Lower lip bearing multiple cones. Outer
plate of maxilla 1 with 7 spines, no spine especially
thickened. Inner plates of maxilliped especially
thin, poorly armed. Coxae 2-4 without special
anterodorsal humps. Some posterior spines on arti-
cle 6 of pereopods 1-2 thin and seta-like, others
thick and stiff, midapical spine or seta absent. Arti-
cle 2 of pereopod 5 without facial setae. Peduncle
of uropod 1 with dorsolateral spines confined api-
488 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
cally or widely spread, medial spines confined api-
cally or widely spread; peduncle of uropod 2 with
only one medial spine confined apically or spines
widely spread. Peduncle of uropod 3 lacking extra
subapical setae or spines. Telson with ordinary pair
of middorsal setules on each side, highly medial.
Gills present on pereonites 2-7 but gills on pe-
reonites 2 and 7 often extremely small, vestigial or
absent, brood plates present only on pereonites 3-5.
Attributes eliminated from descriptions of spe-
cies: posteriormost seta (minute) on coxae 1-3; fa-
cial ridge formula on article 2 of pereopods 3-5 =
0-1-1.
TYPE-SPECIES.?Limnoporeia kingi Fearn-Wan-
nan, 1968 [monotypy].
COMPOSITION.?Following new species: Limno-
poreia kalduke, L. maranowe, L. ungamale, L. wak-
kine, L. woorake, and L. yarrague.
RELATIONSHIP.?Limnoporeia bears certain re-
semblances to the Brolgus group of genera in terms
of its rounded-classification epimeron 3 and combed
peduncle of uropods 1-2, but otherwise, Limno-
poreia differs in its fully chelate gnathopods, uni-
articulate palp of maxilla 1 and in the odd rnan-
dibular molar bearing shortened to obsolete spines
with development of a fuzzy spine or apical plaque.
Apparently Limnoporeia belongs with the Meta-
phoxus group of genera, by virtue of similar max-
illae, maxillipeds, epimera, uropods, enlarged and/
or poorly setose gnathopods. Limnoporeia differs
from Metaphoxus primarily in the fully chelate
gnathopods.
The species are arranged in the order of increas-
ing complexity of the telson. This is associated
with an increasing complexity of setae on pereo-
pods 1-2, and, to some extent, in the increasingly
stronger development of gills on gnathopod 2 and
pereopod 5.
Key to the Species of Limnoporeia and Vldanamia
1. Telson with only one spine on each lobe besides any setules 2
Telson with 2 or more spines on each lobe besides any setules 6
2. Epimeron 2 with all setae arranged in tandem horizontally S
Epimeron 2 with 2-3 setae arranged in vertical row 5
3. Article 2 of antenna 1 elongate, dactyls of pereopods 1-2 ordinary', bearing inner acclivity
L. woorake, new species
Article 2 of antenna 1 not elongate, dactyls of pereopods 1-2 elongate, lacking inner acclivity . 4
4. Outer ramus of uropod 1 naked, epimeron 2 with 3 facial setae, epimeron 4 with 4 posterior
setules, eyes small and dark, oramatidia small, article 6 of pereopod 4 with one posterior
seta, coxae 1-4 with 5 4 1 1 setae narrowly confined L. kingi
Outer ramus of uropod 1 with one dorsal spine, epimeron 2 with 2 facial setae, epimeron 4
with 3 posterior setules, eyes large, clear, ommatidia enlarged, article 6 of pereopod 4 with
3 posterior setae, coxae 1-4 with 8-9-9-4 setae widely spread L. yarrague, new species
5. Dactyls of pereopods 1-2 partially reduced in size, telson ic lobes with apicomedial protrusion
JL maranowe, new species
Dactyls of pereopods 1-2 of normal phoxocephalid size, telsonic apices with apicolateral pro-
trusion L. wakkine, new species
6. Dactyls of pereopods 1-2 vestigial, discoid, telson with two sets of apical spines on each lobe,
one set subdistal, ventral apex of rostrum with fully conical and sharp protrusion
V. friUare, new species
Dactyls of pereopods elongate, unguiform, telson with only one set of apical spines on each
lobe, ventral apex of rostrum with rudimentary or no protrusion 7
7. Epimeron 2 with pair of vertically oriented setae almost fully posterior, dactyls of pereopods
1-2 ordinary, each telsonic lobe with 2 spines, outer ramus of uropod 2 with 0-1 spine
L. ungamale, new species
Epimeron 2 with pair of vertically oriented setae set near middle of epimeron, dactyls of
pereopods 1-2 elongate, curved, each telsonic lobe with 3 spines (one or more thin), outer
ramus of uropod 2 with 3 spines L. kalduke, new species
NUMBER 245 489
Limnoporeia kingi Fearn-Wannan
Limnoporeia kingi Fearn-Wannan, 1968:3&-43, figs. 4-7.
HOLOTVPE.?NMV J.153, female, 3.20 mm [newly
measured].
TYPE-LOCALITY.?Jones Bay, Lake King, Victoria,
Australia, May 1957.
VOUCHER MATERIAL.?Type-locality, 4 paratypes.
REMARKS.?The holotype is very similar to L.
yarrague as described below, with pereopods 3 and
5 similar to that species and not as illustrated by
Fearn-Wannan, figs. 4B.C; article 2 of pereopod 5
is somewhat more protuberant posteriorly than in
L. yarrague; article 6 of pereopod 4 bears 'only one
posterior seta. The eyes are dark, smaller than
those of L. yarrague, and the ommatidia are con-
cealed, smaller and more irregularly sized and dis-
tributed. Apical spines on the telson are shorter
than in L. yarrague (despite Fearn-Wannan's fig-
ure) and the setae on coxae 1-4 are fewer, = 5-4-
4-1, not counting anterior setule of coxae 1-3 nor
2 short setules of coxae 4. Epimera 2 and 3 are out
of phase with uropod 1 as epimeron 2 has 3-4 fa-
cial setae, epimeron 3 has 4 posterior setules while
the outer ramus of uropod 1 lacks a dorsal spine.
In L. yarrague epimeron 2 has 2 setae, epimeron 3
has 3 setules while the outer ramus of uropod 1
has one dorsal spine. The most proximal group of
spines on article 4 of antenna 2 in L. kingi is com-
posed of 3 spines, 2 of these of normal length, the
third extremely elongate.
Adults of L. yarrague reach a larger size than
those of L. kingi as far as known and some of the
paucities of L. kingi may be phenotypic owing to
earlier maturation, but the nonsymmetric situation
among epimera and uropod 1 suggest that L. kingi
and L. yarrague are distinct species.
In respect of head, antennae 1-2, gnathopods,
pereopods, especially dactyls of pereopods 1-2, epi-
mera, uropods, and telson, the holotype is ex-
tremely close to the illustrations herein presented
of L. yarrague. The holotype was not dissected to
obtain views of mouthparts nor telson; the telson
of L. kingi appears to be somewhat shorter than
that of L. yarrague, although Fearn-Wannan shows
it with elongate lobes and spines nearly as long as
those of L. yarrague; each lobe apically bears one
spine and one lateral setule. Fearn-Wannan's illus-
trations show the palp of maxilla 1 of L. kingi to
be shorter than in L. yarrague and article 4 of
antenna 2 lacking the third group of spines but
the holotype bears those 2 spines and a setule, for
a formula of 3-3-2; L. kingi also bears, according
to Fearn-Wannan's figures, a seta on article 2 of
the mandibular palp, absent in Western Port L.
yarrague.
Limnoporeia yarrague has also been discovered
in Lake King sympatric with L. kingi. One addi-
tional specimen of L. kingi was found in a collec-
tion of 207 specimens, 206 of which are L. yarra-
gue. The Lake King specimens of L. yarrague are
minutely distinct from L. yarrague in Western Port
in that article 2 on the mandibular palp bears 2
subapical setae and thus resembles L. kingi in this
character. The ventral setae on coxae 1^ 4 are fewer
and variable in number (5, 6, 7, 8), unlike L. yar-
rague in Western Port. However, L. yarrague from
Lake King bears the usual 2 ventral setae on epi-
meron 2 and 2-3 posterior setules on epimeron 3.
A sample from Merimbula, New South Wales, also
contains L. kingi sparsely and L. yarrague in great
abundance.
Paratypes of L. kingi have been examined as
follows: (1) ovigerous female is L. kingi; (2) sub-
adult male with early proliferation of flagellum on
antenna 2 is L. kingi; (3) a large headless specimen
with excessive epimeral setation appears to be L.
kingi on the basis of subtle generalities in urosome
and pereopods; (4) a subadult female is L. yarra-
gue. Each specimen has at least one subapical seta
on article 2 of the mandibular palp so that in Lake
King, L. yarrague and L. kingi are inseparable on
this character difference found in Western Port L.
yarrague. One additional difference between the
two species in Lake King is the presence of 8 setal
spines apically on article 3 of the mandibular palp
in L. kingi and 10 spines in L. yarrague.
MATERIAL.?NMV, one sample (4, fifth specimen
referrable to L. yarrague). Resample of type-
locality, Lake King, Paynesville (1). MMD, one
sample, 10 Feb 1972 (2).
DISTRIBUTION.?Victoria, Lake King. New South
Wales, Merimbula, intertidal, on edge of channel.
Limnoporeia maranowe, new species
FIGURES 247-249
DESCRIPTION OF FEMALE.?Head about 17 per-
cent of total body length, greatest width about 65
490 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s
FIGURE 247.?Limnoporeia maranowe, new species, female "d," 2.77 mm (m
2.86 mm).
male "m,
NUMBER 245 491
percent of length; rostrum long, reaching at least
to apex of article 2 on antenna 1, constricted
weakly towards apex (from dorsal view), apex
somewhat attenuate but rounded. Eyes small, com-
posed of black core in preservative surrounded by
occasional small and clear ommatidium. Article 1
on peduncle of antenna 1 about 1.7 times as long
as wide, almost twice as wide as article 2, apico-
ventral region with 4 setules, strongly produced
dorsal apex with about 4 setae; article 2 about 0.6
times as long as article 1, ventral apex with 4 setae;
flagella extending subequally, primary and acces-
sory flagella each 4-articulate, aesthetascs absent or
one alone present on accessory fiagcllum. Article 3
of antenna 2 with one long, one short facial setae;
spine formula of article 4 = 3-3-2, proximal group
thin, both proximal groups with accessory setule,
dorsal margin with weak notch bearing spine and
setae, ventral margin with 2 groups of 2-3 setae
and 3 subbasal penicillate setules, ventrodistal
margin with 3 short to medium spines; article 5
almost 0.7 times as long as article 4, facial spine
formula = 1 plus accessory setule, dorsal and ven-
tral margins straight and smooth, dorsodistal apex
with 2 spines, ventrodistal apex with 2 spines and
2 setae; flagellum as long as article 5 of peduncle,
3-articulate. Prebuccal complex convex anteriorly,
moderately extended forward, epistome and upper
lip articulate, epistomal part small, upper lip with
widely spread lateral alae, ventral margin bisinuate
from anterior view. Mandibles with weak palpar
hump; right incisor with 3 teeth and notch; left
incisor with 5 humps in 2 branches; right lacinia
mobilis broadly flabellate, minutely and complexly
toothed and bearing facial tooth; left lacinia
mobilis broadly flabellate, with 5 teeth; right rakers
2, left 3; mandibular molar small, projecting, sub-
conical, heavily setulose, these setules attached to
strongly cornified, clear cone on apex of molar;
palp article 2 bearing one inner seta subapically,
article 3 about 0.85 times as long as article 2,
strongly oblique apex with 8 setae. Lower lip with
4 cones on outer lobe, one cone facial and disjunct
from others, mandibular lobes of medium length,
rounded apically, inner lobes small, fused together
for most of their length. Inner plate of maxilla 1
large, elliptical, naked; outer plate with 7 spines,
two of these with one cusp each on opposite sides;
palp exceeding apex of outer plate, uniarticulate,
thin, bearing 2 apical and one subapical setae.
Inner plate of maxilla 2 broader and slightly
shorter than outer plate, setae sparse, inner and
outer plates each with one medial seta. Inner plate
of maxilliped with one apical and one medial seta;
outer plate with 4 medial spines; palp article 2
sparsely setose medially, article 3 with 2 facial
setae, article 4 of medium length, not fully ungui-
form, bearing one apical spine and 2 accessory
setae. Coxa 1 unexpanded distally, setal formula of
coxae 1-4 = 5-5-5-2, anterior and posterior mar-
gins of coxa 4 parallel, posterodorsal excavation
medium, V-shaped, ratio of width to length = 5:6.
Gnathopod 1 especially slender, gnathopod 2 as
long as but stouter than gnathopod 1; long pos-
terior setae on article 2 of gnathopods 1-2 = 1 and
1, small anterior setae on both members = one
middle, one distal; article 3 of both gnathopods,
but especially of gnathopod 2, slightly elongate;
article 5 on gnathopod 1 long, with flat and free
posterior margin, shorter on gnathopod 2, posterior
margin somewhat triangulate and almost cryptic;
article 6 of both gnathopods long, slightly tapering
apically, strongly chelate, apex of palm bearing
scale-like protrusion and medium spine, apices of
dactyls with especially large apical scale; length
ratios of articles 5-6 on gnathopods 1-2 = 69:112
and 55:132, width ratios = 24:36 and 2T:49.
Pereopods 1-2 generally similar to each other but
articles 4-5 of pereopod 2 slightly stouter than on
pereopod 1; long posterior setae on article 2 of
pereopods 1-2 = 3 and 4, anterior short setae = 2
and 1, articles 4-5 lacking facial armaments; article
4 with 2 groups of sparse simple posterior setae,
produced anterodistal apex with 2 setae; article 5
with several posterior setae, one long distal spine
reaching to M. 80 on article 6; article 6 with one
midposterior seta and apical rows of 3 and 2 spines
on pereopod 1, 3 and 3 on pereopod 2, spines
mostly long; dactyl extremely short, curved, coni-
cal, lacking acclivities, nail absent (unless repre-
sented by chitinous internal remnants). Coxae 5-7
posteroventral setule formula = 2-1-3. Articles 4-5
of pereopods 3-4 narrow, facial spines almost ab-
sent; width ratios of articles 2, 4, 5, 6 of pereo-
pod 3 = 52:23:18:10, of pereopod 4 = 68:20:15:9,
of pereopod 5 = 91:16:13:7, length ratios of pe-
reopod 3 = 68:26:29:30, of pereopod 4 = 90:50:
39:42, of pereopod 5 = 100:17:27:22; article 2
of pereopod 5 extending to apex of article 4;
medial apex of article 6 combed, lacking digits.
492 SMITHSONIAN CONTRIBUTIONS TO ZOOLOG1
FIGURE 248.?Limnoporeia maranowe, new species, female "d," 277 mm (m
2.86 mm).
male "m,
NUMBER 245 493
Posteroventral corners of epiraera 1,2,3, =
rounded, rounded-quadrate, and quadrate, pos-
terior margins = flat, weakly convex, and almost
flat, with 3 setules on epimeron 3, epimeron 2
with 5 facial setae, anteriormost 2 of these hori-
zontal, posteriormost 3 of these vertical. Urosomite
1 naked ventrally. Apicolateral corner on peduncle
of uropod 1 smooth, with small spine, medial apex
with long spine, medial margin with small spine in
middle; each ramus lacking apical nail but with
weak scale or tiny apical setule, each ramus with
one small dorsal spine, apicolateral corner on
peduncle of uropod 2 smooth, with one medium
sized apicolateral spine, one smaller apicomedial
spine, rami like those of uropod 1 apically, outer
ramus with dorsal spine, inner naked. Uropod 3 of
medium length, peduncle with one ventrolateral
spine, one dorsal setule; rami feminine, inner ex-
tending to M. 60 on article 1 of outer ramus, inner
ramus broad basally, very slender apically, tapering
strongly near base, ramus appearing curved basally,
bearing one short apical seta, article 1 of outer
ramus bearing 2 lateral spines, one apicolateral
spine, one long apicomedial spine-seta, article 2
elongate, 0.37, bearing 2 apical setae. Length-width
ratio of telson about 3:2, not fully cleft, each apex
narrow but with narrow medial lobe, broad lateral
acclivity bearing one short spine and one lateral
setule of similar length, middorsal pair of plu-
setules diverse. Cuticle bearing very sparse but un-
usual bulbar setules, each setule with short tube-
like internal tail (nerve duct?), twin bulb and ex-
tremely long spike-like external seta and very short
accessory setule. Gills on gnathopod 2 and pereo-
pod 5 vestigial.
DESCRIPTION OF MALE.?Following parts like fe-
male: head, prebuccal and upper lip, left mandible,
lower lip, maxillae, maxilliped, generalities of
gnathopods and pereopods 1-2. Eyes enlarged. An-
tenna 1 with medial patch of fuzz (stout setules)
on article 1; primary flagellum with 5 articles, first
2 each bearing calceolus, basal articles and article
3 of peduncle with stout aesthetascs. Spines and
setae on peduncle of antenna 2 shorter than in
female, articles 3-4 with dorsomedial fuzz, article 4
facial formula = 3-2-2; article 5 of peduncle rela-
tively longer than in female, ventral margin con-
vex, dorsal margin with one dish-shaped calceolus
and 2 sets of male setules; flagellum elongate, bear-
ing 33-35 articles, lacking calceoli. Teeth and facial
cusp of right lacinia mobilis much sharper than in
female; article 2 of palp with 2 inner apical setae,
article 3 with 9 apical setae. Coxae 1-3 each with
only 3 long ventral setae, coxa 4 with only one
long ventral seta. Long posterior setae on article
2 of gnathopods 1-2 and pereopods 1-2 = 1-1-2-4,
short anterior setae = 2-1-2-1. Article 6 of gnatho-
pod 1 slightly more elongate than in female. Arti-
cle 6 of pereopods 1-2 each with spine rows of 3
and 3. Article 2 of pereopods 3-5 slightly to
greatly narrower than in female, pereopod 4 rela-
tively longer than in female. Epimera 1-2 lacking
facial ridges, epimeron 2 with only one facial seta,
epimeron 3 with 2 posterior setules. Urosomite 1
bearing setule at base of uropod 1; urosomite 3
with stronger dorsal protrusion basally than in
female. Spines on uropods 1-3 very small, apical
scales of rami slightly larger than in female. Rami
of uropod 3 masculine, inner ramus extending al-
most to apex of article 1 on outer ramus, medial
margins of inner ramus and outer ramus setose,
outer margin of inner ramus naked. Telson slightly
more elongate than in female, apical spine slightly
smaller than in female, rudimentary denticle row
present on each lobe near proximal end of deft.
Gill present on pereopod 5.
OBSERVATIONS (male).?The poorly developed
calceoli, aesthetascs, and denticle rows on the telson
suggest that the males at hand are not fully ma-
ture.
ILLUSTRATIONS.?Male coxa 1 magnified same di-
mensions as male coxa 4 after latter balanced to
female coxa 4; male coxae 1-4 thus of similar pro-
portions to those of female.
HOLOTYPE.?NMV, female "a," 3.06 mm.
TYPE-LOCALITY.?CPBS 42S, 1 Nov 1971, West-
ern Port, Victoria, Australia, 15.1 m, sand.
VOUCHER MATERIAL.?Type-locality: male "m,"
2.86 mm (illus.); female "d," 2.77 mm (illus.); male
"c," 3.15 mm.
RELATIONSHIP.?Because this species is used as
a model for the genus, its relationships are dis-
cussed in the comparisons made for other species.
Limnoporeia maranowe is selected as the model be-
cause of its regular phoxocephalid telson in com-
bination with ordinary head, fully developed right
lacinia mobilis, and elongate apical setae on the
outer ramus of uropod 3. However, L. maranowe
is not the perfect model of primitiveness because
of its alate upper lip and shortened dactyls of
494 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 249.?I.imnoporeia maranowe, new species, female "d," 2.77 mm (TO
2.86 mm).
male "m,
NUMBER 245 495
pereopods 1-2.
MATERIAL.?CPBS, 23 samples from 15 stations
(41); PPBES, 3 samples from one station (5).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, 8-19 m, sand, muddy sand, sand on
clay.
Limnoporeia yarrague, new species
FIGURES 250-254
DESCRIPTION OF FEMALE.?Head about 18 per-
cent of total body length, greatest width slightly
less than 60 percent of length; rostrum long, reach-
ing past apex of article 2 on antenna I, uncon-
strictcd, tapering evenly, apex rounded. Eyes med-
ium-small, ochraccous, free of dark pigment in
preservative. Article 1 on peduncle of antenna 1
about 1.5 times as long as wide, almost twice as
wide as article 2, apiconventral region with about
5 setules (all not illustrated), strongly produced
dorsal apex with 4 setules; article 2 about half as
long as article 1, ventral apex with 3 setae; acces-
sory flagellum much shorter than primary flagel-
lum, 3-articulate, primary 5-articulate, aesthetascs
present, bases tubular, apices flat. Article 3 of an-
tenna 2 with one long, one short facial setae; spine
formula of article 4 = 3-3-2, proximal group thin,
both proximal groups with accessory setule, dorsal
margin with weak notch bearing spine and seta,
ventral margin with 2 groups of 3-4 setae and 3
subbasal pen kill ate setules, ventrodistal margin
with 3 medium to long spines, article 5 almost 0.6
times as long as article 4, facial spine formula = 1
plus accessory setule, dorsal margin straight and
smooth, ventral margin with one acclivity bearing
long seta and 2 setules, dorsodistal apex with 3
medium to long spines, ventrodistal apex with one
spine, setule and long seta; flagellum as long as
article 4 of peduncle, 4-articulate. Prebuccal com-
plex flat anteriorly, poorly extended forward, flush
with rnandibular margins, poorly lobed dorsally,
epistome and upper lip fused together but regions
distinguishable, upper lip not widely alate, ventral
margin broadly rounded, anterior face with cres-
centic ridge. Mandibles with strong palpar hump;
right incisor with 3 teeth, notch and accessory
tooth; left incisor with 5 humps in 2 branches;
right lacinia mobilis broadly flabellate, poorly de-
veloped, short, bearing one facial hump, biting
margin weakly sinuate; left lacinia mobilis broadly
flabellate, with 6 irregular teeth; right rakers 2, left
3; mandibular molar small, projecting, heavily
setulose, these setules attached to strongly cornified
clear cone on apex of molar; palp article 2 naked,
article 3 about 0.8 times as long as article 2,
strongly oblique apex with 10 setae. Lower lip with
small contiguous pair of cones on each outer lobe,
mandibular lobes long, broad, asymmetrically
rounded apically, inner lobes broad, fully fused
together and bearing longitudinal facial hump,
apex of fused lobes with unusually long setules.
Inner plate of maxilla 1 large, subovate, naked;
outer plate with 7 spines, two of these with 1-2
cusps each on opposite sides, no spine especially
thickened; palp exceeding apex of outer plate,
thin, bearing 3 apically flush setae. Inner plate of
maxilla 2 broader than and almost flush with outer
plate, setae sparse but long, inner plate with med-
ial seta on back face submarginally; outer plate
with 2 medial setae. Inner plate of maxilliped long,
narrow, with 2 apical setae; outer plate long and
thin, with 3 medial spines; palp article 2 moder-
ately setose medially (double rows), article 3 with
2 facial setae, article 4 of medium length, almost
fully unguiform, bearing one long apical spine and
2 accessory setae. Coxa 1 weakly expanded distally;
setal formula of coxae 1-4 = 8-(9-10)-9-4; coxa 4
very large, anterior and posterior margins weakly
divergent, posterodorsal excavation medium, al-
most L-shaped, ratio of width to length = 15:16.
Gnathopod 1 slender, gnathopod 2 as long as but
stouter than gnathopod 1; long posterior setae on
article 2 of gnathopods 1-2 = 1 and 3, small anter-
ior setae on both members = 1 and 2; article 5 on
gnathopod 1 long, with produced but then flat-
tened posterior margin, this article short on
gnathopod 2 and almost cryptic; article 6 of both
gnathopods long, constricted in middle, curving
and expanding apically, strongly chelate, apex of
palm bearing scale-like protrusion and medium
spine; apices of dactyls curved but simple and
sharp, article 3 of both gnathopods weakly and
equally elongate; length ratios of articles 5-6 on
gnathopods 1-2 = 65:110 and 55:116, width ra-
tios = 28:43 and 29:52 (base of article 6 = 35
and 47). Pereopods 1-2 generally similar to each
other; long posterior setae on article 2 of pereo-
pods 1-2 = 5 and 6, each with one short posterior
seta, naked anteriorly, articles 4-5 lacking facial
armaments except for one apicodorsad seta on arti-
496 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s
FIGURE 250.?Limnoporeia yarrague, new species, holotype, female "a," 3.86 mm (Y ? aesthetasc).
cle 5, article 4 with several groups of weakly den-
dritic posterior setae, produced dorsal apex with
one long, one short setae, article 5 with several
posterior setae, one long distal spine reaching
about to M. 75 on article 6; article 6 with one mid-
posterior seta on pereopod 1, joined by spine on
pereopod 2, article 6 with apical rows of 3 and 3
spines (or setae); dactyl very long, deeply hooked
and sharp, lacking acclivities and nail and basal
outer setule. Coxae 5-7 posteroventral setule for-
mula = 0-0-0. Articles 4-5 of pereopods 3-4 nar-
row, facial spines absent; width ratios of articles
NUMBER 245
497
FIGURE 251.?Limnoporeia yarrague, new species, holotype, female "a," 3.86 mm.
498 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Rl
FIGURE 252.?Limnoporeia yarrague, new species, holotype, female "a," 3.86 mm.
2, 4, 5, 6 of pereopod 3 = 68:22:19:9, of pereo-
pod 4 = 81:26:17:9, of pereopod 5 = 95:23:17:8,
length ratios of pereopod 3 = 88:29:26:32, of
pereopod 4 = 105:59:59:70, of pereopod 5 = 120:
24:29:20; article 2 of pereopod 5 extending to
apex of article 4; medial apex of article 6 sinu-
ate and combed. Posteroventral corners of epimera
1, 2, 3 = rounded, quadrate, and weakly protuber-
ant, posterior margins = almost flat, weakly con-
vex, and weakly convex, with one setule each on
epimera 1-2 and 3 on epimeron 3, epimeron 2 with
2 facial setae, lacking ridge. Urosomite 1 naked
ventrally. Apicolateral corner of peduncle on uro-
pod 1 with medium spine and strong combs, med-
ial apex with long spine, margins otherwise naked,
each ramus of uropods 1-2 with poorly defined
apical nail (largely obsolescent) and scale, latter
poorly developed on uropod 2, rami of uropod 1
each with dorsal spine, rami of uropod 2 naked
dorsally, peduncle of uropod 2 with one apicolat-
eral, one apicomedial spine, apicolateral corners
with combs. Uropod 3 long, peduncle with 2 ven-
trolateral spines, one apicomedial spine, rami
feminine, inner extending to M. 60 on article 1 of
outer ramus, inner ramus lanceolate, bearing one
vestigial setule, article 1 of outer ramus attenuate
apically, bearing 2 lateral spines, one apicolateral
spine, distance between spines 2 and 3 much
greater than between spines 1 and 2, article 2 elon-
gate, 0.40, bearing 2 short apical setae. Length-
width ratio of telson about 19:13, not fully cleft,
each apex very narrow, truncate, bearing long
spine and short lateral setule, middorsal pair of
short plusetules diverse. Cuticle bearing very sparse
but ordinary bulbar setules with very short setular
part emerging into shallow dishpits, epimera with
very fine and weak striations. Gill of gnathopod 2
absent, of pereopod 5 vestigial.
OBSERVATIONS (specimens from Merimbula,
N.S.W.).?Smallest ovigerous females available.
Female "I," 2.81 mm: Mandibular palp article
2 with one inner apical seta, article 3 with 8. Spine
formula on article 4 of antenna 2 = 3-3-1. Setal
formula of coxae 1-4 = 6-7-7-1. Article 6 of pereo-
pods 1-2 with 2 disjunct posterior setal-spines.
Article 2 of pereopod 5 with only 4 posterior teeth.
Epimeron 1 lacking ridge; epimeron 3 with only
one posterior setule notch. Article 1 on outer
ramus of uropod 3 with only one lateral spine
acclivity, article 2 almost 60 percent as long as
article 1.
Male "m," 3.15 mm: Eyes enlarged, mostly oc-
cluded by pigment. Article 1 of antenna 1 with
medial patch of aesthetascs (fuzz); article 2 with 3
apicoventral setae; accessory flagellum 3-articulate;
primary flagellum 5-articulate, both bearing
aesthetascs. Articles 3-4 of antenna 2 with dorso-
medial patches of fuzz, articles 4-5 stout, spine
formula on article 4 = 3-3-1 or 3-3-2, ventral
margin with 4 short apical setae and setules, apico-
distal margin with 3 short spines, article 5 spine
formula = 1, apicoventral margin with small spine
and several setules, midapex with one spine, dorsal
apex with 2 spines and calceolus, flagellum elon-
gate, flagellar formula = 38, 2, 3, 5, 7 . . . 21 or 2, 4,
6, . . . 20. Article 2 of mandibular palp with 3 long
NUMBER 245 499
mXl
FIGURE 25S.?Limnoporeia yarrague, new species (m ? male "m," 3.15 mm; K ? seta of pereo-
pods 1-2).
inner apical setae, article 3 with 10 apical setae;
form of right lacinia mobilis shown in illustration.
Inner lobes of lower lip more tumid than in female
(see illustration). Inner plate of maxilla 1 more cir-
cular than in female (see illustration). Outer plate
of maxilla 2 with 6 setae. Coxa 1 relatively smaller
in ratio to coxa 4 than in female, shape of latter
like that of female; setal formula on coxae 1-4 =
6-8-8-3. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 1-1-2-3,
short anteriors = 2-2-1-0, no others; article 6 of
gnathopod 1 slightly stouter than in female, ratio
of lengths of articles 5-6 of gnathopod 1 =64:102,
widths = 22:36. Article 6 of pereopods 1-2 with
spine formula of 3-3 plus 2 setal spines disjunct on
posterior margin. Pereopod 3 smaller relative to
pereopod 5 than in female, article 2 of pereopod
4 thin. Epimeron 2 slightly enlarged, epimeron 3
shortened posteriorly, lengthened anteriorly (see
illustration), epimeron 1 lacking ridge and setae,
epimeron 2 with 2 setae, no ridge, epimeron 3 with
2 posterior setule notches. Medial margin of
500 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 254.?Limnoporeia yarrague, new species (m ? male "m," 3.15 mm; / ? female "f."
2.81 mm).
peduncle on uropod 1 with 2 spines. Peduncle of
uropod 3 with 2 ventral spines; rami submasculine,
inner reaching almost to apex of article 1 on outer
ramus, setose medially, with one lateral seta and 2
apical setae, article 1 of outer ramus with one
lateral acclivity, spine formula = 1-1, setal formula
= 1-1, article 2 about 42 percent as long as article
1, bearing 2 short apical setules. Telson elongate,
ratio of length to width = 44:27, apices narrowly
truncate, each bearing lateral setule, one medium
spine and one medial longer spine, dorsal denticle
rows absent. Urosome highly modified; urosomite 1
with large dorsal crest-hump; urosomite 2 forming
saddle between urosomites 1 and 3; urosomite 3
with erect dorsal hump. Like female: gnathopods
1-2, pereopods 1-2, coxae 1?4 except for items spe-
cifically mentioned.
OBSERVATIONS (specimens from EBS).?Male "d,"
3.66 mm, EBS 138: Epimeron 2 without setae;
flagellum on antenna 2 with 46 articles; humps on
urosomites 1 and 3 more strongly produced than
in figured male; setae of pereopods 1-2 as in super-
female described below.
Senile superfemale "e," 5.25 mm, EBS 103:
Head elongate. Eyes enlarged as in male. Flagellum
of antenna 2 with 9 articles, basal articles short, ap-
pearing .in proliferative phase. Setal formula of
coxae 1-4 = 10-12-10-6. Epimeron 3 with 2 pos-
terior notches. Uropods 1-2 and outer ramus of
uropod 3 as in holotype, inner ramus of uropod 3
elongate, reaching to about M. 80 along article 1
of outer ramus, not marginally setose. Telson as in
figured male, with 2 apical spines and setule on
each lobe. Urosomite 1 with rudimentary hump
but urosomite 3 unproduced dorsally. Apices of
setae on pereopods 1-2 with trigger extension rela-
tively smaller and shorter than in small adults and
not secondarily branched. Broodplates of pereo-
nites 3-5 fully developed.
Specimens from EBS 103, 304, and 313 are some-
what smaller than the holotype but several are sex-
ually mature; female "g," 3.36 mm, from EBS 313
bears 4 eggs. The eyes of all specimens are darkly
occluded with purple pigment except for periph-
eral fractions of a few ommatidia (specimens held
in formaldehyde only for short time). In other re-
spects these specimens fit the holotype, except for
female "g," which has an elongate article 2 on the
NUMBER 245 501
outer ramus of uropod 3, this article being 51 per-
cent as long as article 1 compared with 42 percent
for the holotype; all others of these dark-eyed speci-
mens have the second article on the outer ramus of
uropod 3 close to the condition figured for the
holotype.
ILLUSTRATIONS.?Outer lobes of lower lip unflat-
tened.
HOLOTYPE.?NMV, female "a," 3.86 mm.
TYPE-LOCALITY.?<JPBS 00E/4, 5 Apr 1965, West-
ern Port, Victoria, Australia, 2.1 m, bottom un-
known.
VOUCHER MATERIAL.?CPBS 00E/2: female "b,"
3.20 nun. MMD, on edge of channel, 10 Feb 1972:
female "f," 2.81 mm (illus.); male "m," 3.15 mm
(illus). F.BS 103: female "e," 5.25 mm. EBS 138:
male "d," 3.66 mm. EBS 313: female "g," 3.36 mm.
EBS 304(1). EBS 103(4).
RELATIONSHIP.?This species resembles Limno-
poreia maranowe in its general appearance but dif-
fers in the following characters, none of which
appears to have a particular association with body
si/e because of equivalent sizes of material studied:
1) the absence of a vertical brush of setae on epime-
ron 2; (2) the elongate dactyls of pereopods 1-2;
(3) the narrow apices and long spines of the telson;
(4) poorly developed right lacinia mobilis; (5) fully
fused inner lobes on the lower lip; (6) poorly alate
upper lip; (7) short accessory flagellum of antenna
1; (8) the ordinary bulbar setules of the cuticle; (9)
presence of apicolateral combs on the peduncles
of uropods 1-2; (10) backside facial seta on inner
plate of maxilla 2. Minor differences include the
shorter apical setae on both rami of uropod 3, the
elongate apical section on article 1 of the outer
ramus of uropod 3, the stouter article 2 of pereopod
3 and much smaller articles 3-7 in relation to
that article 2, pereopod 3 being greatly larger in
relationship to the size of pereopod 5 in this
species than in Limnoporeia maranowe, and
finally, the stouter and more pyriform article 2 of
pereopod 4.
See L. kingi for distinctions from that species
and notes on L. yarragite in Lake King.
MATERIAL.?CPBS, 2 samples (2); MMD, 3 sam-
ples (75); EBS, 4 samples from 4 stations (9); Lake
King, Victoria, Paynesville, one sample (206).
DISTRIBUTION.?Victoria: Western Port, 2 m,
muddy sand; Lake King. New South Wales: Botany
Bay, Towra Beach, 0.9 m, Zostera and shell grit;
Merimbula, intertidal, muddy sand; Georges River,
Woronora, 3 m, sand.
Limnoporeia woorake, new species
FIGURES 255-257
DESCRIPTION OF FEMALE.?Head about 15 percent
of total body length, greatest width slightly more
than 60 percent of length; rostrum long, reaching
about to middle of article 2 on antenna 1 (but this
article unusually elongate), unconstricted, tapering
evenly, apex rounded. Eyes of medium size, ochra-
ceous, free of dark pigment in preservative. Article
1 on peduncle of antenna 1 about 1.35 times as
long as wide, about 1.7 times as wide as article 2,
apicoventral region with about 6 setules, strongly
produced dorsal apex with 2 setules; article 2 about
as long as article 1, ventral apex with 3 long setae;
accessory flagellum slightly shorter than primary
flagellum, accessory flagellum 3-articulate, primary
4-articulate, aesthetascs very weak if not absent.
Article 3 of antenna 2 with one medium, one
short facial setae; spine formula of article 4 = (1?)
-3?4?2, thick formed, both proximal groups with
accessory setule, one spine of most proximal group
elongate and seta-like, dorsal margin with one short
thick spine, ventral margin with 2 penicillate
setules, one apical acclivity with 2 short setae, ven-
trodistal margin with one seta, 2 spines, followed by
one seta; article 5 about 0.8 times as long as article
4, facial spine formula = 3 plus one accessory setule,
dorsal margin straight and smooth, ventral margin
with one acclivity bearing 2 medium stiff setae,
dorsodistal apex with 2 spines, ventrodistal apex
with one long spine and 2 subdistal setae; flagellum
about 0.4 times as long as article 5 of peduncle,
3-articulate. Prebuccal complex flat anteriorly,
poorly extended forward, flush with mandibular
margins, poorly lobed dorsally, epistome and upper
lip fully articulate dorsally, upper lip poorly alate,
anterior margin with weak crescentic rugosity
highly ventrad, lower margin with hump defined
laterally on both sides by weak sinus. Mandibles
with medium-sized palpar hump; right incisor with
3 teeth and ripples, accessory hump in crotch be-
tween 2 proximal teeth; left incisor with 5 humps
in 2 branches; right lacinia mobilis broadly flabel-
late, well developed, with 3 facial teeth, distal mar-
gin variously toothed; left lacinia mobilis broadly
502 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 255.?Limnoporeia woorake, new species, holotype, female "a," 4.75 mm (n ? male "n,
3.57 mm; u ? female "u," 3.76 mm).
NUMBER 245 503
flabellate, with 5 teeth; right rakers 2, left 3;
mandibular molar small, projecting, heavily setulose,
these setules attached to weakly distinct but corni-
fied clear cone occasionally subdivided into 2
branches; palp article 2 with one inner basal setule,
article 3 about 1.1 times as long as article 2, oblique
apex with 4-5 setae. Lower lip with small pair of
cones on each outer lobe, mandibular lobes of ordi-
nary extension, apices narrow and rounded, inner
lobes of ordinary width, fused together except for
deep apical incision, bearing longitudinal facial
hump also divided apically. Inner plate of maxilla
1 large, ovate, naked, attached by broad base (un-
usual); outer plate with 7 spines, none with cusps
on opposite sides; palp reaching only to apex of
outer plate, of medium stoutness, bearing 2 api-
comedial and one apicolateral setae. Inner plate
of maxilla 2 much broader than and flush with
outer, setae on inner plate very short, one seta
medial, setae on outer plate very long, one seta
medial. Inner plate of maxilliped with one apical,
one apicolateral and one medial setae; outer plate
with 4 medial spines; palp article 2 moderately
setose medially (double rows), article 3 with 3
facial setae, article 4 of medium length, almost fully
unguiform, bearing one short apical spine and 2
accessory setae. Coxa 1 strongly expanded distally,
adz-shaped; setal formula of coxae 1-4 = 3-3-2-0;
coxa 1 occasionally with midventral bulbar setule;
anterior and posterior margins of coxa 4 divergent,
with long posterodorsal lobe, posterodorsal excava-
tion medium, almost L-shaped, ratio of width to
length = 41:39. Gnathopod 1 very slender (except
for article 6), gnathopod 2 as long as but much
stouter than gnathopod 1, long posterior setae on
article 2 of gnathopods 1-2 = 0 and 2, short setae
= 0 and 3, anterior margins with 4 and 2 short
setae, article 3 of both gnathopods of normal length;
article 5 of gnathopod 1 highly elongate, with
weakly extended posterior margin, article 5 of
gnathopod 2 shorter and almost cryptic; article 6
of gnathopods large but diverse, both weakly
chelate, apex of palm on gnathopod 2 bearing scale-
like protrusion and medium spine, scale thin and
spine-like, gnathopod 1 lacking scale but with
strong scale-like elongate tooth and small spine,
article 6 of gnathopod 1 expanding distally, pos-
terior margin undulant, this article on gnathopod 2
of more regular trapezoidal form, apex of dactyl on
gnathopod 1 subflagellate. Length ratios of articles
5-6 on gnathopods 1-2 = 92:74 and 59:90, width
ratios = 21:55 and 20:66. Pereopods 1-2 generally
similar to each other; long posterior setae on article
2 of pereopods 1-2 = 7 and 6, long anterior setae
= 1 and 0, short anterior setae = 2 and 1; article 4
slightly shorter on pereopod 2, articles 4-5 lacking
facial armaments, posterior margin of article 4
with 2 groups each of 4-5 and 2-3 simple setae,
article 5 with several posterior setae and one un-
usual, short, apically flagellate spine, long distal
spine reaching about to M. 60 on article 6; article
6 with one short stout midposterior spine, apical
rows of 3 and 1 spines, dactyl of normal phoxo-
cephalid proportions, with strong inner acclivity in
form of hook bearing contiguous long setule, apical
nail absent or invisible, base of dactyl with ordi-
nary long plusetule. Coxae 5-7 posteroventral setule
formula = 1-1-1. Articles 4-5 of pereopods 3-4 of
medium stoutness, facial spines moderately devel-
oped anteriorly; width ratios of articles 2, 4, 5, 6
of pereopod 3 = 60:27:21:12, of pereopod 4 =
78:28:20:11, of pereopod 5 = 115:22:20:10, length
ratios of pereopod 3 = 70:23:30:28, of pereo-
pod 4 = 90:45:39:28, of pereopod 5 = 112:27:
31:18; article 2 of pereopod 5 slightly exceed-
ing apex of article 4; medial apex of article 6
slightly ragged but not combed. Posteroventral
corners on epimera 1,2, 3, = rounded, rounded,
protuberant and bearing setule sinus, posterior
margins all protuberant, with one setule each on
epimera 1-2, one seta and one setule on epimeron
3 in deep notches between protuberances, epimeron
2 with one facial seta. Urosomite 1 naked ventrally.
Apicolateral corner of peduncle on uropod 1 with
small spine, no combs, medial apex lacking spine
but margin at base of outer ramus with weak
comb; peduncle of uropod 2 with one apicolateral
spine, medial apex with setule, no combs on uro-
pod 2; rami of uropods 1-2 all naked dorsally,
apical nails large, fused and marked only by stria-
tion bank, each bearing scale, mediobasal margin
of inner ramus on uropod 1 with tiny setule. Uro-
pod 3 long, peduncle with one ventral spine, one
apicolateral and one apicomedial spine; rami
feminine, inner ramus very short, leaf-like, apex
bearing setule, reaching about M. 40 on article 1 of
outer ramus, article 1 of outer ramus bearing one
lateral acclivity armed with one short stout spine
and 2 long setae, similar armament apicolaterally,
medial apex with 2 setae, article 2 slender and ordi-
504
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 256.?Limnoporeia woorake, new species, holotype, female "a," 4.75 mm (n = male "n,"
3.57 mm).
nary, 0.23, bearing 2 apical setae. Length-width
ratio of telson about 6:5, not fully cleft, each apex
of ordinary width, weakly concave, armed with
medium spine and one larger lateral setule, mid-
dorsal pair of medium plusetules subequal. Cuticle
bearing very sparse and unusually tiny bulbar
setules barely emerging into wide and shallow dish-
pits, epimera with very weak and fine striations.
Gnathopod 2 with small, leaf-like gill, pereopod 5
with vestigial, knob-like gill.
DESCRIPTION OF MALE.?Head about 19 percent
of total body length, eyes enlarged, with deep pur-
ple core in preservative surrounded by 1.0?1.5 lay-
ers of ommatidia. Primary flagellum of antenna 1
NUMBER 245
505
nW
FIGURE 257.?Limnoporeia woorake, new species, holotype, female "a," 4.75 mm (n =? male "n,"
S.57 mm).
506 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
with 5-6 articles, aesthetascs much stronger than
in female, primary flagellum with one calceolus
each on articles 1 and 2, articles 2-3 otherwise like
those of female, ventromedial face of article 1 of
peduncle bearing patch of fuzz, elements very stout.
Articles 3-4 of antenna 2 like those of female but
dorsal margins with fuzz composed of very stout
elements almost of magnitude of aesthetascs, dorsal
spine of article 4 absent, article 5 broader than in
female, facial spine formula = 2, ventral margin
with 2 sets of short setae and setule, dorsal apex
with only one small spine; flagellum elongate,
about one third of body length, flagellar formula
= (16-18), 1-3, 5, 7 . . . (15 or 17). Mandibular palp
article 3 with 7 apical spine-setae. Coxae 1 and 3
like those of female, coxa 2 about 10 percent taller
than in female, coxa 4 slightly narrower (antero-
posterior) than in female, coxa 2 with only 2 pos-
teroventral setae. Long posterior setae on article 2
of gnat ho pods 1-2 and pereopods 1-2 = 1-2-4-4,
anterior short setae = 1-0-1-1, medium anterior
setae = 0-1-0-0, long anterior setae = 0-0-1-0.
Article 4 of pereopods 1-2 slightly thinner than in
female and bearing posterior groups of only 4 and
1 setae or 4 and 2. Pereopod 3 much smaller and
pereopod 4 slightly smaller relative to pereopod 5
than in female; article 2 of pereopod 3 much nar-
rower, of pereopod 4 slightly narrower than in
female, article 2 of pereopod 5 narrower than in
female. Epimera wider than in female, posterior
margin of epimeron 3 generally with only one long
seta and one setule at corner but young males with
epimeron 3 like that of female. Urosomite 1 with
dorsal denticles. Peduncle of uropod 2 with 2 api-
codistal spines, medial apices on peduncles of uro-
pods 1-2 at base of outer ramus with combs, strong
and few members on uropod 1. Uropod 3 mascu-
line, strongly setose, article 1 of outer ramus with
2 lateral acclivities each armed with one short
spine and 2 long setae, inner ramus extending only
to M. 70 on article 1 of outer ramus, peduncle
lacking apicolateral spine. Telson longer than in
female, main apical spine on each lobe shorter than
in female and apicolateral plusetule longer, each
lobe with basomedial row of dorsal denticles. Gill
on pereopod 5 well developed.
VARIATIONS.?Female "f," 3.66 mm: article 5 of
antenna 2 with only 2 facial spines but epimeron 4
with 4 posterior notches and article 1 of uropod
3 with 2 sets of lateral spine-setae. Male "g," 3.43
mm: epimeron 3 with 5 notches posteriorly and
article 5 of antenna 2 with stout posterior spine
(and setule) in place of thin spine (and setule) seen
in figured male of this species.
HOLOTYPE.?NMV, female "a." 4.75 mm.
TYPE-LOCALITY.?CPBS A3 5, 6 Aug 1964, West-
em Port, Victoria, Australia, 15.4 m, bottom un-
known.
VOUCHER MATERIAL.?PPBES 986/5: male "n."
3.57 mm (illus.); female "u," 3.76 mm (illus.); male
"d," 3.43 mm; plus 3 other females and one male.
CPBS 32E: female "f," 3.66 mm. RHM: male "g,"
3.43 mm, and one other male.
RELATIONSHIP.?This species can be distinguished
from both Limnoporeia maranowe and /.. yarrague
in the presence of only one facial seta on epimeron
2, in the ordinary dactyls of pereopods 1-2 bearing
the normal acclivity and setule, in the elongate
article 2 of antenna 1, and in the apices of the
telson which arc broader than those of /.. yarrague
and lack the medial lobe of /.. maranowe. The
inner plate of maxilla 2 is much broader than in
the other 2 species, and maxilla 1 differs in: the
broad basal attachment of the inner plate, the short
and stouter palp, and the absence of opposite
cuspidation on certain spines of the outer plate.
Mandibular palp article 2 is somewhat elongate in
this species. The normal right lacinia mobilis
stands in contrast to that of L. yarrague. The stout
spines of antenna 2 are reminiscent of species
thought to be inquilinous in Syndexamine and
even Kondoleus. Proportions of gnathopods also
distinguish this species from others (see illustra-
tions).
MATERIAL.?CPBS, 3 samples from 3 stations (3);
RHM (4); PPBES, 5 samples from 3 stations (27).
DISTRIBUTION.?Victoria: Western Port and Port
Phillip Bay, neritic and 4-15.4 m, sand, sandy
gravel.
Limnoporeia ungamale, new species
FICURES 258-260
DESCRIPTION OF FEMALE.?Head about 20 percent
of total body length, greatest width about 50 per-
cent of length; rostrum very long, exceeding apex
of article 2 on antenna 1, unconstricted, apex nar-
row but rounded, apicoventral face of rostrum
weakly thickened. Eyes of medium size, circular,
NUMBER 245
507
FIGURE 258.?Limnoporeia ungamale, new species, holotype, female "a," 4.29 mm (n; -= female
"w," 3.62 mm).
508 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
composed of black core in preservative surrounded
by one layer of clear ommatidia. Article 1 on
peduncle of antenna 1 about twice as long as wide,
twice as wide as article 2, apicoventral region with
8 setules, strongly produced dorsal apex with 2
setules; article 2 about 0.5 times as long as article I,
ventral apex with 4-5 setae; accessory flagellum
slightly shorter than primary flagellum, latter with
4 articles, former with 3-4 articles, aesthetascs ab-
sent or perhaps one alone present. Article 3 of an-
tenna 2 with one long, one short facial setae; spine
formula of article 4 = 3-3-5 or 3-3-3, proximal
spines thin, both proximal groups with accessory
setules, dorsal margin with weak notch bearing
spine and seta, ventral margin straight and naked
except for subbasal penicillate setules and one mid-
marginal setule, ventrodistal margin with 2 med-
ium to short spines and group of 3 setal spines;
article 5 almost 0.7 times as long as article 4, facial
spine formula = either 1 or 2 plus accessory setule,
dorsal margin straight and smooth, ventral margin
with acclivity bearing 2-3 spines plus setule, dorso-
distal apex with 2 spines, seta and setule, mid- and
ventrodistal apex with 2 long spine-setae, 2 setules;
flagellum as long as article 5 of peduncle, 4-articu-
late. Prebuccal complex with epistomal hump mod-
erately extended forward, epistome and upper lip
fused together, upper lip with rounded but ex-
tended lateral humps, ventral margin with weak
middle hump, weak anterior crecentic ridge.
Mandibles with strong palpar hump; right incisor
with 3 teeth, notches and ripples; left incisor with
5 humps in 2 branches; right lacinia mobilis
broadly flabellate, minutely and complexly toothed,
bearing toothed facial hump and several small
facial teeth; left lacinia mobilis broadly flabellate,
with 5 teeth; right rakers 2, left 3; mandibular
molar small, projecting, subconical but apically
divided into 4 spiniform and setulose teeth; palp
article 2 with one subbasal short inner seta, article
3 about 1.3 times as long as article 2, strongly
oblique apex with 8 setae. Lower lip with 2-3
cones on outer lobe, mandibular lobes of medium
length, rounded apically, inner lobes broad, fused
together for most of their length but oral and
aboral faces bearing two separate humps or one
fused plaque. Inner plate of maxilla 1 large, sub-
ovate, naked, base very broad; outer plate with 7
spines, one of these with one cusp on opposite side;
palp exceeding apex of outer plate, medium-thin,
bearing 5 apical and subapical setae, one subapical
seta in medial notch. Inner plate of maxilla 2 ex-
tremely broad and slightly shorter than outer
plate, setae sparse, one small medial seta; outer
plate with several medial setae. Inner plate of
maxilliped with 2 apical and one medial setae;
outer plate with 3 medial spines; palp article 2
moderately setose medially (2 rows), article 3 with
4 facial setae, article 4 of medium length but bear-
ing long apical nail and almost fully unguiform,
accessory setae on apex of article 4 either 1 or 2,
second seta if present elongate. Coxa 1 expanded
apically; setal formula of coxae 1-4 = (5-6)-(5-7)-
(5-7)-(3); coxa 4 strongly quadratiform, anterior
and posterior margins parallel, |>osterodorsal exca-
vation shallow but wide. V-shaped, ratio of width
to length = 11:12. Gnathopod 1 csj>ct tally slender;
gnathopod 2 as long as but stouter than guathopod
1; long posterior setae on article 2 of gnathopods
1-2 = 1 and 1, short anterior setae 3 and 3; arti-
cle 3 of both gnathopods slightly elongate; article
5 on gnathopod 1 long, with Hat and free posterior
margin, this article much shorter on gnathopod 2,
posterior margin somewhat triangular and almost
cryptic; article 6 of both gnathopods long, tapering
slightly on gnathopod 2 but constricted in middle
on gnathopod 1 and reexpanding apically, strongly
chelate apices of palms and curved apices of dactyls
similar on both sets of gnathopods, each with long
thin scale-like extension and medium spine, apical
scale on dactyls scarcely distinct, pointed, essen-
tially nail-like; length ratios of articles 5-6 on
gnathopods 1-2 = 87:97 and 60:127, width ra-
tios = 21:38 and 28:54. Pereopods 1-2 generally
similar to each other; long posterior setae on arti-
cle 2 of pereopods 1-2 = 2 and 4, short setae =
1 and 0, short anterior setae = 5 and 4, only pe-
reopod 1 with posterodistal corner bearing 2 setae
(not in normal posterior formula), strongly shifted
proximally; articles 4-5 lacking facial armaments,
article 4 with 3 groups of dendritic posterior setae,
produced anterodistal apex with seta and setule,
article 5 with dense posterior setae, long distal
spine reaching almost to M. 80 on article 6; article
6 with 2 midposterior sets of 3 setae each (rarely
2 setae in set), apical and posterior rows of 4 and 3
spines, spines mostly long and medium; dactyl of
medium (normal) length, almost straight, lacking
acclivities and setules but weakly constricted api-
cally and bearing partially fused blunt apical nail.
NUMBER 245
509
FIGURE 259.?Limnoporeia ungamalc, new species, holotype, female "a," 4.29 mm.
510
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Rll
FIGURE 260.?Limnoporeia ungamale, new species, holotype, female "a." 4.29 mm (w - female
"w," 3.62 mm).
Coxae 5-7 posteroventral setule formula = 1-1-1.
Articles 4-5 of pereopods 3-4 narrow; facial spines
almost absent or weakly developed on anterior
margins of articles 4-6 of pereopod 3; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 64:21:15:9,
of pereopod 4 = 80:27:16:10, of pereopod 5 =
102:17:12:8, length ratios of pereopod 3 = 80:
23:30:33, of pereopod 4 = 97:67:49:47, of pereo-
pod 5 = 115:24:33:25; article 2 of pereopod 5
reaching apex of article 4; medial apex of arti-
cle 6 on pereopod 5 indistinctly ragged. Postero-
ventral corners on epimera 1, 2, 3 = rounded,
rounded-quadrate, and weakly protuberant, poste-
rior margins = protuberant, convex, and convex,
with one setule each on epimera 1 and 3, epimeron
2 with one midfacial seta and 2 vertical setae at
posteroventral corner. Urosomite 1 naked ventrally
but with seta at lateral base of uropod 1; urosomite
2 with similar but smaller seta. Apicolateral corner
of peduncle on uropod 1 smooth, with small spine,
midlateral margin with 0-1 small spine, medial apex
with medium spine, medial margin with 2 other
smaller spines, rami of uropods 1-2 with apical nail
obsolescent but weak scale present, outer ramus of
uropod 1 with one dorsal spine, inner with 1-2,
outer ramus of uropod 2 with 0-1 spine, inner
naked, comb present only on medial apex of uropod
2 peduncle at base of outer ramus, medial margin
of uropod 1 peduncle minutely fuzzy but not
combed. Uropod 3 highly elongate, peduncle with 2
ventrolateral spines, dorsally with small medial
spine, lateral setule, rami feminine, inner extending
to M. 35 on article 1 of outer ramus, inner ramus
leaf-like, tapering evenly, bearing apical setule, arti-
cle 1 of outer ramus with 4 lateral acclivities each
with short spine and long seta, similar set apicolater-
ally, one spine apicomedially, article 2 elongate,
bearing 2 apical setae and marginal setules. Telson
of ordinary length, length-width ratio = 33:28, not
fully cleft, each apex broad, weakly excavate but
basically truncate and inwardly oblique, bearing me-
dium medial spine, small lateral spine and lateral-
most setule, middorsal pair of plusetules diverse.
Cuticle with numerous bulbar setules and weak
striations. Gnathopod 2 with large gill, pereopod
5 with small but perceptible gill.
ILLUSTRATIONS.?Palpar hump of mandible simi-
lar to that shown for L. yarrague.
HOLOTYPE.?NMV, female "a," 4.29 mm.
TYPE-LOCALITY.?CPBS 51N, 15 Jan 1972, West-
ern Port, Victoria, Australia, 16.5 m, sand and
mud.
VOUCHER MATERIAL.?CPBS C6/1, female "w?"
3.62 mm (illus.). Male unknown.
NUMBER 245 511
RELATIONSHIP.?The species is unique in the
genus for the presence of 2 stout apical spines on
each apex of the telson. The dactyls of pereopods
1-2 are of ordinary length, neither elongate nor
stunted, lack acclivities or setules and have a poorly
denned apical nail-spine. The pair of vertically
oriented posterior setae of epimeron 2 are very
close to the corner, and this feature combined with
the presence of only one ventrofacial seta is an-
other good mark of identity. The head is very
elongate but the apicoventral protrusion on the
rostrum is very weak. Pereopods 1-2 have complex
setae. All of these characters distinguish this spe-
cies from Limnoporeia maranowe, L. yarrague, and
L. woorake.
Limnoporeia ungamale has the unusual kind of
inner plate on maxilla 1 seen in L. woorake but
the gnathopods of these two species are so strik-
ingly distinct that there can only be an indirect
relationship between the two species. See L. kal-
duke for further distinctions.
MATERIAL.?CPBS, 2 samples from 2 stations (2).
DISTRIBUTION.?Victoria, Western Port, 10-16.5
m, sand, muddy sand.
Limnoporeia wakkinr, new species
FIGURES 261-263
DESCRIPTION OF FEMALE.?Female "b": Head
about 20 percent of total body length, greatest
width about 40 percent of length; rostrum very
long, reaching end of article 2 on antenna 1, un-
constricted or weakly so apically, apex narrow but
rounded, apicoventral face of rostrum weakly thick-
ened. Eyes of medium size, subcircular, composed
of black core in preservative surrounded by occa-
sional clear ommatidia. Article 1 on peduncle of
antenna 1 about 2.1 times as long as wide, twice
as wide as article 2, apicoventral region with about
7 setules, strongly produced dorsal apex with 4
long setules; article 2 about 0.53 times as long as
article 1, ventral apex with 5 setae; accessory fla-
gellum 4-articulate, slightly shorter than primary
ftagellum, latter also 4-articulate, one aesthetasc
present apically on each flagellum. Article 3 of
antenna 2 with one medium-long, one short facial
setae; spine formula of article 4 = 3-4-4, proximal
spines thin, both proximal groups with accessory
setule, dorsal margin with weak notch bearing
spine, ventral margin straight and naked except for
medial penicillate setules extending unusually dis-
tad, ventrodistal margin with 3 medium to long
spines and group of 3 short setae; article 5 about
0.75 times as long as article 4, facial spine for-
mula = 1 plus accessory setule, dorsal margin
straight and smooth, ventral margin with acclivity
bearing 2 spines plus seta and setule, dorsodistal
apex with 3 short to long spines and several setae,
ventrodistal apex with long spine-seta and 2 set-
ules; flagellum as long as article 5 of peduncle,
3-articulate, basal article elongate. Prebuccal com-
plex weakly undulant anteriorly, moderately ex-
tended forward, epistome and upper lip fused to-
gether, upper lip with weakly excavate ventral
margin, weak anterior crescentic ridge. Mandibles
with strong palpar hump; right incisor with 3 teeth,
notches and ripples; left incisor with 7 humps in 2
branches; right lacinia mobilis broadly flabellate,
bifid, proximal branch resembling raker spine with
papillate setules, distal branch weakly serrate dis-
tally, one facial hump near base of proximal
branch; left lacinia mobilis broadly flabellate, with
5 teeth; right rakers 2, left 3; mandibular molar
large, forming plaque with apex weakly projecting
and bearing 3-4 fixed spine-teeth and one com-
plexly penicillate short seta; palp article 2 with one
subbasal short inner seta, article 3 about 1.15 times
as long as article 2, strongly oblique apex with 8
spine-setae. Lower lip with 4 cones on outer lobe, 2
of these facial, mandibular lobes short, almost sub-
acute apically, inner lobes of moderate size, fused
together for most of their length but aboral face
bearing plaque. Inner plate of maxilla 1 large, sub-
ovate, naked, base broad; outer plate with 7 spines,
none of these with cusp on opposite side; palp
exceeding apex of outer plate, medium-thin, bear-
ing 5 apical and subapical setae, most subapical
setae in weak medial notch. Inner plate of maxilla
2 extremely broad and slightly shorter than outer
plate, setae sparse, one medial seta and 2 large facial
setae; outer plate with several medial setae. Inner
plate of maxilliped with attenuate apex bearing one
seta, apicolateral acclivity bearing second seta,
medial margin with seta; outer plate with 3 medial
spines; palp article 2 strongly setose medially (3
rows), article 3 with 4 facial setae, article 4 elon-
gate, bearing medium apical nail, two accessory
setae, one highly disjunct. Coxa 1 expanded dis-
tally; setal formula of coxae 1-4 = 6-7-8-3; coxa
512 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 261.?Limnoporeia wakkine, new species, female "b," 4.24 mm (Y = rostrum).
NUMBER 245 513
4 strongly quadratiform, anterior and posterior
margins parallel, posterodorsal excavation shallow
but wide, V-shaped, ratio of width to length = 1:1.
Gnathopod 1 especially slender, gnathopod 2 as
long as but slightly stouter than gnathopod 1; long
posterior setae on article 2 of gnathopods 1-2 =
1-1, short anterior setae = 3-4, no others; article 3
of both gnathopods slightly elongate but more so
on gnathopod 2 (length ratio of this article on
gnathopods 1-2 = 35:45); article 5 on gnathopod
I long, with flat and free posterior margin, this
article shorter on gnathopod 2, posterior margin
bluntly triangular but not cryptic; article 6 of both
gnathopods long, on gnathopod 2 tapering slightly
but on gnathopod 1 weakly constricted in middle
and reexpanding apically; strongly chelate apices of
palms and curved apices of dactyls similar on both
sets of gnathopods, each with long thin scale-like
extension and medium spine, apical scale on dactyls
scarcely distinct, pointed, essentially nail-like;
length ratios of articles 5-6 on gnathopods 1-2 =
87:90 and 66:115, width ratios = 18:34 and
27:47. Pereopods 1-2 generally similar to each
other and to L. ungamale but articles 5-6 slightly
smaller; long posterior setae on article 2 of pereo-
pods 1-2 = 2 and 3, short anterior setae = 5 and 5,
pereopod I with normal distal placement of apico-
posterior setal pair on article 2; articles 4-5 lacking
facial armaments, article 4 with 3-4 groups of sim-
ple posterior setae (1 or 2 to 6 setae per group),
produced anterodistal apex with 1-2 setae (one a
setule), article 5 with dense posterior setae, long
distal spine reaching almost to M. 80 on article 6;
article 6 with 2 midposterior sets of 3 setae each,
rarely 4 (one pereopod only), apical and posterior
rows of 4 and 3 spines, spines mostly long and
medium; dactyl of medium (normal) length, almost
straight, lacking acclivities and setules and bearing
almost fully invisible apical immersed nail. Coxae
5-7 posteroventral setule formula = 1-1-4. [Articles
6-7 of pereopod 3 and 5-7 of pereopod 4 missing];
articles 4-5 of pereopods 3^4 narrow, facial spines
almost absent or weakly developed on anterior
margin of articles 4-5 of pereopod 3; width ratios
of articles 2, 4, 5, 6 of pereopod 3 = 65:20:16:?,
of pereopod 4 = 80:25:?:?, of pereopod 5 = 100:
21:13:9, length ratios of pereopod 3 = 78:24:
28:?, of pereopod 4 = 91:65:?:?, of pereopod
5 = 110:24:35:25; article 2 of pereopod 5 exceed-
ing apex of article 4. Posteroventral corners
on epimera 1, 2, 3, = rounded (to weakly-rounded-
quadrate), rounded-quadrate, and weakly but
sharply protuberant, posterior margins all convex,
with one setule on each epimeron, epimeron 2 with
4 facial setae, anteriormost 2 of these horizontal,
posterior 2 of these set vertically (broken in illustra-
tion). Apicolateral corner of peduncle on uropod 1
smooth, with spine (as long as in L. ungamale) and
one midlateral similar spine, medial margin un-
known, damaged, rami of uropods 1-2 with apical
nail, obsolescent but weak scale present, outer
ram us of uropod 1 with 2 dorsal spines twice as long
as those on L. ungamale, inner with 2 of similar di-
mensions, rami of uropod 2 naked, peduncle of
uropod 2 with 2 lateral spines of same size illus-
trated for L. ungamale, ventrolateral margin of
peduncle on uropod I with 3 short setae, combs
absent on uropodal peduncles. (Uropod 3 mostly
missing, unknown.) Length-width ratio of telson =
6:5, not fully cleft, each apex broad, weakly exca-
vate but basically truncate and inwardly oblique,
bearing long medial spine-seta, 0-1 small lateral
seta and lateralmost setule, middorsal pair of
plusetules weakly diverse, elongate. Urosomite 1
naked ventrally but with small seta at lateral base of
uropod 1; urosomite 2 with similar seta. Cuticle
with bulbar setules and weak striations. Gnathopod
2 with small, leaf-like gill, pereopod 5 with vestigial,
knob-like gill.
Female "a," 3.70 mm: Differences from female
"b": eyes fully clear of pigment; ventral apex of
article 2 on antenna 1 with 4 setae, primary flagel-
lum with aesthetasc also on articles 2-3; spine
formula on article 4 of antenna 2 = 4-4-4, article
5 with ventral notch bearing 2 long spinules and
long setule; mandibular molar with only 2 spine-
teeth, palpar apex with 7 spine-setae; palp of
maxilla 1 with only 4 setae; long setal formula (in-
cluding small posterior seta on coxae 1-3) of coxae
1_4 = (6-7)-(7)-(7-8)-(3-4); long posterior setae on
article 2 of gnathopod 2 = 1, of pereopods 1-2 = 3
and 4, small anterior setae on article 2 of pereopods
1_2 = 3 and 3; article 4 of pereopods 1-2 with only
3 groups of 3-5 posterior setae; setal formula of
coxae 5-7 = 1-1-2; width ratios of articles 2, 4, 5,
6 of pereopod 3 = 61:18:15:8, of pereopod 4 =
79:28:15:8, of pereopod 5 = 95:17:13:8, length
ratios of pereopod 3 = 75:25:27:30, of pereopod
4 = 87:59:45:39, of pereopod 5 = 102:20:30:21;
epimeron 2 with 2 ventral setae, posterior face
514 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 262.?Limnopcreia wakkine, new species, holotype, male "h," 4.72 mm (a - female "a,"
3.70 mm).
NUMBER 245 515
with only one seta, epimeron 3 with or without
midposterior setule notch; medial margin of
peduncle on uropod 1 with 3 large spines, outer
ramus of uropod 1 with only one dorsal spine,
outer ramus of uropod 2 with one dorsal spine;
peduncle of uropod 3 with 3 ventral spines, dorso-
laterally with one spinule, medially with one spine,
rami feminine, inner short and scale-like, reaching
to M. 48 on article I of outer ramus, with one tiny
apical setule, article 2 of outer ramus elongate, 0.40,
bearing 2 long apical setae, apicomedial corner of
article 1 with one long spine, lateral margin with 2
acclivities, spine formula = 1-2-2, setal formula =
0; length-width ratio of telson = 15:11, each apex
truncatoconcave, hearing tiny lateral setule, then
larger setule, then very elongate setal-spine, then
shorter setal-spine and finally with small medial
setule.
OBSERVATIONS (female).?Female "a" differs from
female "b" primarily in the elongation of the spines
in the ventral notch on article 5 of antenna 2, in
the presence of only one posterior facial seta on
epimeron 2 and in the much heavier apical spino-
setation on each lobe of the telson.
Four females accompanying female "a," all speci-
mens of slightly smaller size, also have the stated
spines on antenna 2 elongate and thin (either hav-
ing 2 or 3) and have the normal telsonic spinoseta-
tion but like female "b" and unlike female "a"
have a pair of posterofacial setae on epimeron 2.
These four, like female "a," have clear eyes. The
outer ramus of uropod 1 bears either 1 or 2 spines,
the inner ramus bears 2, the outer ramus of uropod
2 bears 0-1 spine, the inner ramus lacks spines. The
spine formula on article 4 of antenna 2 varies in
these ways: 3-3-3 and 3-3-2. Long setae on coxae
1-4 vary in these ways: 6-5-6-3, 6-5-5-3, 5-5-5-3,
5-5-5-2. Both flagella of antenna 1 bear 4 articles.
DESCRIPTION OF MALE (male "h," holotype).?
Head about 19 percent as long as body, eyes slightly
enlarged, mainly obscured by pigment. Mandibles,
lower lip, maxillae 1-2 as described for female "b"
but see illustrations of right lacinia mobilis and
mandibular palp. Article 1 of antenna 1 with num-
erous apicoventral setules; accessory flagellum
4-articulate; primary flagellum 5-articulate, articles
1-4 bearing large medial aesthetascs, article 1 with
rows of 3, 7, and 11 aesthetascs, article 2 with 11
apical, article 3 with 8 apical, and article 4 with one
apical aesthetascs (some omitted in illustration);
medial face of article 1 on peduncle with small
patch of fuzz. Articles 3-4 of antenna 2 with medio-
dorsal fuzz, article 3 with dorsodistal spine, spine
formula on article 4 = 3-4-4; apical and ventral
spines on article 5 thin, flagellum elongate, 17-
articulate, calceoli absent (male young as evidenced
by smallness of medial patch of fuzz on antennae
1-2 and shortness of inner ramus on uropod 3). No
spine on maxilla 1 with opposite cuspidation. Nail
on article 4 of maxillipedal palp shorter than in
female. Setae on coxae 1-4 = (7)-(7)-(7)-(4-5); coxa
4 narrower than in female, ratio of length to
width = 19:16. Long posterior setae on article 2 of
gnathopods 1-2 and pereopods 1-2 = 1-1-3-3,
short anteriors = 2-3-3-3, no others. Pereopods 1-2
alike; pereopods 1-4 generally like those of female,
pereopod 5 larger; ratios of widths on articles
2, 4, 5, 6 of pereopod 3 = 59:17:15:8, of pereo-
pod 4 = 75:26:15:8, of pereopod 5 = 100:15:13:7,
ratios of lengths of pereopod 3 = 77:21:28:30,
of pereopod 4 = 88:60:45:40, of pereopod 5 =
107:21:31:21; medial apex of article 6 on pereo-
pod 5 finely combed. Epimera 1-2 lacking lateral
ridge, epimeron 2 with only one small facial seta.
Peduncle of uropod 1 with 4 medial spines, of uro-
pod 2 with one large apicomedial spine, lateral
margin of peduncle on uropod 1 with 1?2 spines,
apex with 1, peduncle of uropod 2 with either 3 or
4 dorsal spines. Rami of uropod 3 submasculine,
inner reaching to M. 70 on article 1 of outer ramus,
apex with 2 setae, medial margin setose, lateral
margin with one seta, article 2 of outer ramus
elongate, bearing 2 medium and short apical setae,
lateral margin of article 1 with 2 acclivities, spine
formula = 1-2-1, setal formula = 1-1-2, medial
margin setose. Telson elongate, ratio of length to
width = 8:5, each apex with 5 elements as in
female "a" but elements shorter. Pipes and bulbar
setules of cuticle much sparser than in female.
OBSERVATIONS.?Male "h" and female "b" with 3
coarse serrations apicoventrally on peduncle of uro-
pod 2, uropod I with fine comb apicoventrally,
male otherwise smooth, female additionally with
fine apicolateral comb on uropod 1, smooth on
uropod 2.
ILLUSTRATIONS.?Many features of this species are
so similar to those shown for L. ungamale that
duplicate illustrations have been omitted, particu-
larly those for the following list of parts: antenna
1, maxilla 1, outer plate and article 3 of maxilliped,
516 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
FIGURE 263.?Limnoporeia wakkine, new species, holotype, male "h," 4.72 mm (a -? female "a,"
3.70 mm).
NUMBER 245 517
gnathopod 1, generalities and especially dactyls of
pereopods 1-2. Details, however, may vary slightly
as given in the description.
Ridge on epimeron 2 of Figure 263: W is ecdysis
line.
Articles 1-5 of gnathopod 2 larger relative to
article 6 than in L. ungamale, but gnathopods
smaller overall than in L. ungamale; anterior spines
on article 2 of pereopod 4 longer than in L.
ungamale (see illustration); article 2 of pereopod 5
slightly broader and longer relative to pereopod 5
of L. ungamale (see dimensions in description;
images of pereopod 5 from both species superim-
posed on each other by matching length of anterior
margin on article 2).
HOI.OTYPK.?VVAM, male "h," 4.72 mm.
TYPE-LOCALITY.?WAM 416-73, trawl sta 6, 24
Jul 1943, Jibbon Point, off Cronulla, New South
Wales, Australia, depth and bottom unknown.
VOUCHER MATERIAL.?Type-locality: female "f,"
2.81 mm. CPBS 51N/172: female "b," 4.24 mm
(illus.). WAM 115-73: female "a," 3.70 mm (illus.);
female "c," 3.95 mm; female "d," 3.82 mm; female
"e," 3.91 mm; female " j , " 4.12 mm.
REMARKS.?The young male is selected as holo-
type because the two adult females, female "b" and
female "a," each have aberrancies as noted in their
descriptions and observations. If any splitting of
this material becomes necessary in the future the
likelihood is that female "b" is a species distinct
from L. wakkine, the latter based on specimens with
thin ventral spines on article 5 of antenna 2.
RELATIONSHIP.?This species is an apparent sib-
ling of L. ungamale. It has far too many minor dif-
ferences from that species to be considered a simple
variant. Fifteen of these distinctions are as follows:
the presence of only one spine in the anterior notch
of article 4 on antenna 2; the considerably distinct
right lacinia mobilis; the essentially unlobate an-
terior lobe of the epistome (lateral aspect); the ab-
sence of opposite cuspidation on any spine of max-
illa 1; the large facial setae on the inner plate
of maxilla 2; the attenuate apex on the inner
plate of the maxilliped; the disjunct accessory
seta on article 4 of the maxillipeds; the broader
coxa 4 and narrower coxa 1 of female; the larger
eye and larger apical protrusion of the rostrum; the
normal placement of the posteroventral setal pair
on article 2 of pereopod 1; the longer posterior lobe
on coxa 6; the lesser posterior position of the ver-
tically set pair of setae on epimeron 2; the longer
spines on the rami of uropod 1; and the elongation
of armaments on the telson.
Limnoporeia wakkine differs from L. kalduke in
the following characters: shorter head, plain upper
lip, in minor details of the right lacinia mobilis, the
disjunct accessory setule on article 4 of the maxil-
lipedal palp, broader apical expansion of coxa 1,
the thin article 5 of gnathopod 1 and elongate arti-
cle 3 of gnathopod 2, the stoutness of the shorter
contiguous spine on article 5 of pereopods 1-2,
the less strongly curved dactyls of pereopods 1-2,
the more posterior position of the setae on epi-
meron 2, the fewer spines on the outer rami of
uropods 1-2 and the nonoblique telsonic apices.
MATERIAL.?CPBS, 2 samples from 2 stations (2);
WAM, 2 samples, 415-73 and 416-73, with only
partial data (10).
DISTRIBUTION.?Victoria, Western Port, 15-16.5
m; New South Wales, off Jibbon Point, 35 m, sand.
Limnoporeia kalduke, new species
FIGURES 264, 265
DESCRIPTION OF FEMALE.?Head about 20 percent
of total body length, greatest width about 40 per-
cent of length; rostrum long, exceeding apex of
peduncle on antenna 1, thin, weakly constricted
near apex, latter attenuate and somewhat acute in
dorsal view but thickened dorsoventrally. Eyes of
medium size, with black core surrounded by occa-
sional small and clear ommatidia. Article 1 of
peduncle on antenna 1 about 2.4 times as long as
wide, about 1.8 times as wide as article 2, ventro-
lateral region with about 7 setules, strongly pro-
duced dorsal apex with 3 long setules; article 2
about 0.3 times as long as article 1, ventral apex
with 5-6 setae; accessory flagellum 5-articulate and
shorter than primary flagellum, latter 6-7 articulate,
one aesthetasc apically on primary flagellum. Arti-
cle 3 of antenna 2 with one long, one short facial
setae, spine formula of article 4 = 3-3-4 or 3-3-5,
spines of proximal group thin, with 3 accessory
setules, middle group with one accessory setule,
dorsal margin with weak notch bearing spine and
seta, ventral margin with one mid group of setae,
one apical group of setae and about 7 subbasal
penicillate setules, ventrodistal margin also with 3
518 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
rNt li,
t^JWi \f/
FIGURE 264.?Limnoporeia kalduke, new species, holotype, female "a," 4.00 mm.
NUMBER 245 519
medium to long spines; article 5 about 0.7 times as
long as article 4, facial spine formula = either 1 or
2 plus accessory setule, dorsal margin straight and
smooth, ventral margin with weak acclivity bearing
3 variously sized setae, dorsodistal apex with 3
spines, ventrodistal apex with 2-3 setae; flagellum
longer than article 4 of peduncle, 5-6-articulate.
Prebuccal complex very flat anteriorly, poorly ex-
tended forward, poorly lobed dorsally, epistome and
upper lip mainly fused but with weak intermittent
articulation ridge, upper lip bell-shaped, ventral
margin with broad, weak, flat, or weakly convex
imagination. Mandibles with strong palpar hump;
right incisor with 3 teeth, accessory tooth, notches
and ripples; left incisor with 6 humps in 2
branches; right lacinia mobilis broadly flabellate,
incipiently bifid, proximal branch spiniform and
armed with papillae as on raker spines, distal
branch broad, minutely serrate distally and with 2
low humps near base of proximal branch; left
lacinia mobilis broadly flabellate, 5-toothed; right
rakers 2, left 3; mandibular molar small, setulose,
poorly projecting, bearing 4-5 teeth, these long and
sharp on left side, all but one on right side small
and blunt; palp article 2 with pair of short setae
near base, article 3 about 1.15 times as long as arti-
cle 2, strongly oblique apex with 8-9 setae. Lower
lip with 4 cones on outer lobe, 2 cones disjunct
from margin, mandibular lobes long, subrounded
apically, inner lobes of medium size, fused together
for most of their length, with pair of facial plaques
in tandem. Inner plate of maxilla 1 large, elliptical,
naked, base somewhat broadened; outer plate with
7 spines, none with opposite cuspidation; palp ex-
ceeding apex of outer plate, thin, bearing marginal
row of 5-6 apical and medial setae. Inner plate of
maxilla 2 broader and shorter than outer plate,
setae sparse, medial face with 2 setae, one enlarged;
outer plate with 3 medial setae. Inner plate of
maxilliped with 2 apical and one medial setae;
outer plate with 3 medial spines; palp article 2
strongly setose medially (3-4 rows), palp article 3
with 4 facial setae, article 4 of medium length,
apical nail of medium length, accessory setae 2.
Coxa 1 scarcely expanded distally, all rounded ante-
roventrally; setal formula of coxae 1-4 = 7-(6-7)-
(6-8)-(3-4); coxa 4 quadratiform, anterior and pos-
terior margins parallel, posterodorsal excavation of
medium size, almost L-shaped, ratio of width to
length = 17:21. Gnathopod 1 especially slender,
gnathopod 2 stouter and longer than gnathopod 1;
long posterior setae of article 2 on gnathopods 1-2
= 1 and 1, small anterior setae on both members
= 2 and 3-4; article 3 of gnathopod 2 slightly elon-
gate; article 5 on gnathopod 1 long, with flat and
free posterior margin, this article shorter on gnatho-
pod 2, posterior margin triangular and almost cryp-
tic; article 6 of both gnathopods long, with taper
followed by weak distal expansion apically, strongly
chelate, apex of palm bearing scale-like protrusion
and medium spine, protrusion blunt; apices of
dactyls ordinary and pointed (presumably scale
fused to dactyl); length ratios of articles 5-6 on
gnathopods 1-2 = 80:103 and 58:135, width ratios
= 25:37 and 30:56. Pereopods 1-2 generally sim-
ilar to each other; long posterior setae on article 2
of pereopods 1-2 = 3 and 4, anterior short setae =
5 and 5-6; article 5 with 1-2 dorsolateral setae on
apical face, article 4 with 3 groups of sparse den-
dritic posterior setae, produced anterodistal apex
with 2 setae, article 5 with dense posterior setae,
long distal spine reaching to M. 75 on article 6;
article 6 with rows of 4 and 2 posterior setae and
rows of 5 and 4 spines apically (same on both pereo-
pods), spines mostly long, dactyl long, sharply
curved, simple. Coxae 5-7 posteroventral setule for-
mula = 1-1-3. Articles 4-5 of pereopods 3-4 nar-
row, facial spines almost absent; width ratios of
articles 2, 4, 5, 6 of pereopod 3 = 63:18:15:8, of
pereopod 4 = 79:24:12:6, of pereopod 5=102:
19:11:6, length ratios of pereopod 3 = 81:22:
27:30, of pereopod 4 = 96:65:51:45, of pereo-
pod 5 = 113:16:30:20, article 2 of pereopod 5
reaching middle of article 5. Posteroventral cor-
ners of epimera 1, 2, 3 = rounded, rounded-quad-
rate, and protuberant with setule sinus, posterior
margins = weakly convex, almost straight, and con-
vex, with one setule each on epimera 1-2, 5 setule
notches on epimeron 3, epimeron 2 with 5 facial
setae, anteriormost 3 of these horizontal, posterior-
most 2 of these set vertically. Urosomite 1 naked
ventrally; urosomites 1-2 each with setule at lateral
base. Apicolateral corner of peduncle on uropod 1
smooth, with small spine, midlateral margin with
one small spine, mediolateral apex with large spine,
medial margin with 2 spines, ventrolateral margin
with several conspicuous setules, rami of uropods
1-2 with apical nails almost fully absorbed, each
with small scale, each ramus of uropod 1 with 3
dorsal spines or outer with only 2, outer ramus of
520 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
FIGURE 265.?Limnoporeia kalduke, new species, holotype, female "a," 4.00 mm.
NUMBER 245 521
uropod 2 with 3?1 dorsal spines, inner ramus with
1-2, peduncle of uropod 2 with one medium apico-
lateral spine and 2 other smaller lateral spines, apex
smooth, medial apex with small spine, medial mar-
gin with smaller spine. Uropod 3 long, peduncle
with 4-5 ventrolateral spines, medial apex with one
spine, dorsolatera! apex with one setule; rami
feminine, inner extending about to M. 63 on article
1 of outer ramus, inner ramus weakly broadened
basally. tapering sharply near apex, bearing 1?2
apical setae, article 1 of outer ramus bearing 3 pairs
of short, subequal lateral spines in acclivities, apico-
latcral coiner with similar pair plus short seta, arti-
cle 2 elongate. 0.10, bearing one short, one medium
apical setae. Length-width ratio of telson about
17:13. not fully cleft, each apex broad, excavate,
oblique, with longer limb lateral, each apex with
subtnaiginal row of 3 long setal spines and one
short lateral setule, dorsomedial pair of plusetules
diverse, highly medial on each lobe. Cuticle finely
striate and bearing small sparse bulbar setules.
Gnathopod 2 with long gill almost reaching end of
article 2, pereopod 5 with small, rounded gill.
OBSERVATIONS.?Female "b," 5.37 mm: Much
larger than holotype, bearing larger eyes (in relation
to head size); inner plate of maxilla 2 with small
facial seta distal to larger (reversed in holotype on
one side only, see illustration of maxilla 2 on holo-
type, presumed aberrant); one lobe of telson with
4 long apical setal spines plus setule; epimeron 2
apparently with posteriormost setal group bearing
3 vertical setae (setae missing, only sockets present).
ILLUSTRATIONS.?View of maxilla 2 aberrant as
smaller facial seta usually set distally from main
facial seta; palmar apices of gnathopods 1?2 similar,
illustrated only for gnathopod 1; except where
described or partially illustrated, view of maxil-
lipeds like that of L. ungamale; in views of dis-
sected uropod 3 and uropod 3 as attached to pleon,
distal two-thirds of outer ramus heavily recon-
structed by approximation as this appendage poorly
preserved and shriveled; setule on coxa 6 missing.
HOLOTYPE.?NMV, female "a," 4.00 mm.
TYPE-LOCALITY.?PPBES 973/3, 12 Feb 1971, Port
Phillip Bay, Victoria, Australia, 13 m, sand. ^
VOUCHER MATERIAL.?Type-locality, female "b,"
5.37 mm; PPBES 973/2, female "c," 4.4 mm. Male
unknown.
RELATIONSHIP.?This species differs from the first
four listed species of this genus in the presence of at
least 3 long spines on each lobe of the telson and in
the presence of 3 (contrast 0-1) spines on the outer
ramus of uropod 2. The elongate dactyls of pereo-
pods 1-2 suggest affinities with L. yarrague but L.
kalduke otherwise differs from that species in the
broad and multispined telsonic lobes, the posterior
vertical set of setae and the presence of a ridge on
epimeron 2. The head of L. kalduke is even longer
and more modified apically than in the first 4
species listed and somewhat transcends those species
and L. wakkine and Uldanamia pillare; see these
latter species for their relationships to L. kalduke.
The vertical set of setae on epimeron 2 suggests
affinities with L. ungamale but that species has
stunted dactyls of pereopods 1-2 and a "normal"
phoxocephalid telson with a medial protrusion on
the apex and one spine and one setule on a lateral
acclivity. Limnoporeia ungamale also has vertically
arranged setae on epimeron 2 but in that species the
setae are set more posteriorly than in L. kalduke,
the telson has only 2 apical spines on each lobe,
both spines being stout and short, and the dactyls
of pereopods 1-2 are of ordinary length, though
plain. Both L. kalduke and L. ungamale share the
same complex setae on pereopods 1-2. From these
various similarities one must suggest that L. kalduke
has its closest affinities with L. ungamale. Neverthe-
less, numerous small differences between these 2
species may be seen. Limnoporeia kalduke is char-
acterized by: (1) more numerous posterior setules
on epimeron 3; (2) shorter setule at base of uropod
1; (3) more numerous ventral setae of epimeron
2; (4) denser armaments of antenna 2; (5) rela-
tively longer inner ramus of uropod 3 on female;
(6) absence of lateral setae on article 1 of uropod 3;
(7) narrower coxa 4; (8) longer coxa 7; (9) stouter
and shorter article 5 of gnathopod 1; (10) normal
placement of posteroventral pair of setae on article
2 of pereopod 1; (11) narrow apex of coxa 1; (12)
absence of opposite cuspidation on any spine of
maxilla 1; (13) differences in upper lip, maxilla 2,
and article 4 on maxillipedal palp, among others.
Even though L. kalduke and L. ungamale appear to
be a close pair of species, the numerous distinctions
between them suggest a manifold and diverse course
of descent from their ancestral stock.
MATERIAL.?PPBES, 4 samples from one station
<6); WPBES, one sample (1).
DISTRIBUTION.?Victoria: Port Phillip Bay and
Western Port, 9-13 m, sand.
522 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCV
Uldanamia, new genus
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae confined apically.
Article 1 of antenna 2 not ensiform; article 3 with
2 setules; facial spines on article 4 in 2 or more
rows, plus special apical spines; article 5 thin. Right
mandibular incisor with 3?4 teeth; molar not tri-
turative, medium, pillow-shaped to elongate?con-
ical, bearing 2 semiarticulate spines and large
plume, bearing fuzz; palpar hump large. Palp of
maxilla 1 uniarticulate; inner plate naked. Setation
of maxilla 2 weak. Inner plate of maxillipeds thin;
apex of palp article 3 not protuberant, dactyl elon-
gate, apical nail distinct, short. Gnathopods similar,
chelate, hands elongate, poorly setose anteriorly;
article 5 free, elongate, with eusirid attachment;
gnathopod 2 weekly enlarged. Article 5 of pereopods
1-2 setose posteroproximally. Article 2 of pereopod
3 of broad form, articles 4-5 of pereopods 3-4 nar-
row to medium, article 2 of pereopods S-A not setose
posteriorly; pereopod 5 ordinary, article 2 naked
ventrally, article 3 ordinary, dactyl very small. Epi-
mera 1-2 lacking long posterior setae, without mid-
facial setae above ventral facial ridge, epimeron 3 of
nonrounded classification and lacking long setae.
Urosomite 1 bearing one or more midventral setae
and setae near base of uropod 1; urosomite 3 with-
out dorsal hook or special process. Peduncle of uro-
pod 1 normally elongate, without apicoventral
spike, without special enlarged apicolateral or
medial spines, peduncular apices of uropods 1-2
not combed, inner ramus of uropod 1 with marginal
spines in one row, no rami continuously spinose to
apex, apical nails weak, inner ramus of uropod 2
ordinary. Uropod 3 ordinary, elongate article 2 of
outer ramus carrying 2 short to medium apical setae.
Telson ordinary, with 6-7 apical spines or setae on
each lobe plus setules, without special dorsal and
lateral spines or setae.
DESCRIPTION.?Rostrum fully developed, with
apicoventral process. Fuzz on article 1 of antenna 1
in male present; calceoli on male primary flagellum
of antenna 1 present. Calceoli on article 5 of male
antenna 2 absent; flagellum in male with calceoli.
Prebuccal parts massive, poorly separated from each
other, upper lip dominant. Right lacinia mobilis
simple; mandibular palp medium to thin, article 1
short, article 2 without outer setae, apex of article
3 oblique. Lower lip bearing multiple cones. Outer
plate of maxilla I with 7 spines, no spine especially
thickened. Inner plates of maxilliped especially
thin, poorly armed. Coxae 2?1 without special an-
terodorsal humps. All posterior spines on article 6
of pereopods 1-2 thin and stiff, midapical spine
absent; dactxls vestigial. Article 2 of pereopod 5
without facial setae. Peduncle of uropod 1 with
dorsolateral spines confined apically, medial spines
widely spread; peduncle of uropod 2 with medial
spines widely spread. Peduncle of uropod 3 lacking
extra subapical setae or spines. Telson with ordi-
nary pair of middorsal setules on each side, highly
medial. Gills present on pereonites 2-7, brood plates
present only on pereonites 3-5.
TYPE-SPKCIES.?Uldanamia pillarc, new species.
COMPOSITION.?Unique.
RELATIONSHIP.? This genus is dearly a product of
the ancestral pool from which Limnoporeia also
descended, and the type-species is simply a highly
specialized limnoporeia in which the following at-
tributes have advanced so far from those found in
Limnoporeia that a separate genus is required to
express the discontiguity.
Uldanamia differs from Limnoporeia in the highly
elongate wrist of gnathopod 2, the special teeth of
epimeron 3 (forcing generic diagnosis to read
"nonrounded classification"), the large size of the
apicoventral process on the rostrum, the vestigial
dactyls of pereopods 1-2, the densely setose pereo-
pods 1-2, and the heavily spinose telson. The state
of most of these characters is more primitive in
Limnoporeia than in Uldanamia.
Uldanamia pillare, new species
FIGURES 266-269
DESCRIPTION OF FEMALE.?Head about 22 percent
of total body length, greatest width about 35 per-
cent of length; rostrum very long, exceeding apex
of peduncle on antenna 1, constricted weakly to-
wards apex (from dorsal view), apex bearing ventral
conical keel-like projection. Eyes small to medium
in size, golden in preservative, clear of pigment core.
Article 1 on peduncle of antenna 1 about twice as
long as wide, almost twice as wide as article 2, api-
coventral region with 8 setules, strongly produced
dorsal apex with 3 setules; article 2 about 0.45 times
as long as article 1, ventral apex with 5-7 setae; ac-
NUMBER 245 523
M
FIGURE 266.?Uldanamia pitlare, new species, holotype, female "a," 4.77 mm (k ? male "k,"
4.82 mm).
524 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Dl
FIGURE 267.?Vldanamia pillare, new species, holotype, female "a," 4.77 mm (k - male "k,
4.82 mm; n ? female "n," 5.04 mm).
cessory flagellum slightly shorter than primary
flagellum, 5-6-articulate, primary flagellum 7-articu-
late, aesthetascs rare or present only apically, articles
elongate. Article 3 of antenna 2 with one long, one
short facial setae; spine formula on article 4 = 4-4-6
or 4-4-7, proximal spines thin, both proximal
groups with accessory setule, dorsal margin with
one spine and 2 setules in group, ventral margin
with group of 4 apical setae plus one seta forming
part of spine group and 6 subbasal penicillate
setules and 2 setules facially, ventrodistal margin
with 5 short to medium spines; article 5 about 0.9
times as long as article 4, facial spine formula = 2
plus accesory setule, dorsal margin straight and
smooth, ventral margin with 2 setal groups, dorso-
distal apex with 2 setae and one spine, ventrodistal
apex with 3 spines, seta and setule; flagellum about
1.4 times as long as article 4 of peduncle, 6-articu-
late. Prebuccal complex convex anteriorly, weakly
extended forward, epistome and upper lip fused
(or epistomal part extremely small if indeed articu-
late, see llustration of male upper lip), ventral mar-
gin of upper lip rounded or weakly excavate (de-
pending on view) and with scarcely perceptible
midhump. Mandibles very bulky and large in rela-
tion to mouthpart field, main body almost cylin-
drical, with incisor and molar projecting from main
body; palpar hump huge; right incisor with 3 teeth,
notches and ripples; left with 6 humps in 2
branches; right lacinia mobilis simple, appearing
similar to raker spine distally, extending onto but
blending basally with molarial body, bearing villi
similar to those of raker spines; left lacinia mobilis
with 5 teeth, right rakers 2, left 3; mandibular
molar forming plaque and extended hump bearing
2-4 teeth, some articulate, plus articulate complex
setule on right molar; palp article 2 naked, article
3 about 1.25 times as long as article 2, oblique apex
with 7-10 spines. Lower lip with 4 cones on outer
lobe, mainly facial (or extremely ill-defined), mandi-
NUMBER 245 525
bular lobes medium to short, rounded apically,
inner lobes broad, fused together for most of their
length but with distinct raphe, with plaque on one
face (inner lobes of maxilla 1 sliding across faces
on inner lobes but edges of plaque forming stop).
Inner plate of maxilla 1 large, elliptical, naked;
outer plate with 7 spines, none with opposite cuspi-
dation; palp exceeding apex of outer plate, thin,
bearing 4 apical and 4 medial setae. Inner plate of
maxilla 2 broader than and extending as far as
outer plate, setae of moderate density, one seta
medial; outer plate with 4 medial setae. Inner plate
of maxilliped very narrow, with 2 medial and 2
apical setae; outer plate with 3 medial spines; palp
article 2 moderately setose medially (2 rows), article
S with 5 facial setae, article 4 long, bearing 2 short
accessory setae, apical spine short. Coxa 1 strongly
expanded distally; coxae 1-4 setal formula = (8-9)-
(9-I0)-10-(3-4), posteriormost seta of coxae 1-3
very short; coxa 4 quadratiform, anterior and pos-
terior margins almost parallel, posterodorsal excava-
tion small, V-shaped, ratio of width to length
= 1:1. Both gnathopods very slender, gnathopod
2 as long as but slightly stouter than gnathopod 1;
long posterior setae on article 2 of gnathopods 1-2
= 1-1, short setae = 2-3, short anterior setae = 3
and 2-3, medium anterior setae = 2-3 and 2; article
3 of both gnathopods elongate; article 5 very long
and narrow, with free and flat posterior margin;
article 6 of both gnathopods of medium length, con-
stricted middistally, then expanding apicalwards,
strongly chelate, apex of palm bearing long blunt
scale-like protrusion and medium spine; dactyl
slightly overlapping palmar margin, apices of
dactyls hooked and with medium to large apical
scale; length ratios of articles 5-6 on gnathopods
1-2 = 120:95 and 129:105, width ratios = 17:27
and 23:36. Pereopods 1-2 especially large, generally
similar to each other but article 4 of pereopod 2
slightly stouter; long posterior setae on article 2 of
pereopods 1-2 = 3 and 4, anterior short setae = 2-4
and 5, articles 4?5 lacking facial armaments except
for 2 anterior setae on article 5, anterodistal corner
of article 4 with 3 setae, posterior margins of articles
4-5 densely setose, in overlapping cycles on article
4, setae dendritic, article 5 with long distal spine
reaching M. 60 on article 6; article 6 with rows of 3
and 1 posterior setae and rows of 5 and 5 spines,
latter short to medium in length; dactyl extremely
short, almost vestigial, crescentic, scarcely pointed,
bearing huge anteromedial setule but otherwise
naked. Coxae 5-7 posteroventral setule formula
= 3-1-3. Articles 4-5 of pereopods 3-4 slightly ex-
panded, facial spines well developed on pereopod 3,
weakly on pereopod 4, facial ridge formula of article
2 = 0?1?1; width ratios of articles 2, 4, 5, 6 of
pereopod 3 = 65:27:24:13, of pereopod 4 = 80:
32:14:9, of pereopod 5 = 109:19:11:6, length
ratios of pereopod 3 = 77=23:32:40, of pereopod
4 = 95:62:41:33, of pereopod 5=110:21:34:23;
article 2 of pereopod 5 almost reaching middle
of article 5. Posteroventral corner of epimeron 1
rounded, of epimeron 2 rounded-quadrate, pos-
terior margins convex, each with setule, epimeron
2 with 5 facial setae, anteriormost 2 of these hori-
zontal, posteriormost 3 of these set obliquely, epi-
meron 3 with sharply protuberant posteroventral
corner, margin above vertical, then turning sharply
oblique and bearing 3 medium upturned teeth ac-
companied by setule in adjacent notch, margin
above teeth almost flat. Urosomite 1 bearing 3 ven-
tral setae on each side and 3-4 anterior setae on
each side. Apicolateral corner on peduncle of
uropod 1 smooth, with small spine, medial apex
with medium to extremely elongate spine (medium
version illustrated), medial margin with 3 small
setae, each ramus of uropods 1-2 with nail almost
fully amalgamated but with strong apical scale,
outer ramus of uropod 1 with 4-6 dorsal spines
inner with 3-4, outer ramus of uropod 2 with 3-4
spines, inner with 1-2, peduncle of uropod 2
smooth, with 4 dorsal spines, medial margin with
2 small spines, one of these apical. Uropod 3 long,
peduncle with 5-7 ventral spines, one mediodorsal
spine; rami feminine, inner slender, extending to
M. 67 on article 1 of outer ramus, bearing one long,
one short apical setae, article 1 of outer ramus with
3-4 lateral acclivities each with 3 spines, one gen-
erally enlarged, one generally small, apicolateral
corner with 3 spines, medial margin setose, article
2 elongate, 0.45, bearing one medium, one short
apical setae. Length-width ratio of telson about
1:1, almost fully cleft, each apex very broad, ir-
regularly truncate, bearing 3 setae and setule, each
apex also with subdistal transverse row of 3 long
spines, middorsal pair of plusetules weakly diverse.
Cuticle with sparse and ordinary bulbar setules, with
faint fingerprint striations especially on basal arti-
cles of pereopods 3-5 and epimera. Gnathopod 2
with long gill almost exceding article 2, pereopod 5
526 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 268.?Vldanamia pillare, new species, holotype, female "a," 4.77 mm (k ? male '%'
4.82 mm).
NUMBER 245
527
W
FIGURE 269.?Uldanamia pillare, new species, holotype, female "a," 4.77 mm (k = male "k,
4.82 mm; Y - aesthetasc).
528 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
with small gill.
DESCRIPTION OF MALE.?Following parts like
female: head; prebuccal: mandibular molars, rakers,
incisors, laciniae mobiles, palpar proportions; lower
lip; maxillae 1-2; maxilliped; gnathopods; pereo-
pods 1-2; coxae 2-3. Eyes enlarged. Antenna 1
bearing dense fuzz on ventral margin of article 1;
primary flagellum 9-articulate, articles short, first 8
each bearing calceolus, first 7 bearing 2-3 large
aesthetascs with tubular bases; accessory flagellum
5-articulate, articles elongate as in female. Articles
4-5 of antenna 2 broader than in female, facial
spines therefore occupying less area, ventral margin
of article 4 with 2 sets of 3 setae, each set obliquely,
plus 2 long marginal setae, facial spines of article 5
longer than in female, dorsal margin with one sub-
distal calceolus, ventral margin with one set of short
setae, ventrodistal corner with short setae, no spines,
dorsodistal corner with 3 long spine-setae; flagellum
highly elongate, articles not elongate, about 47-
articulate, flagellar formula = (?47), 1-5, 8, 10, 12
. . . (44 or 46). Mandibular palp article 2 bearing 2
inner apical setae, apex of article 3 with 8 spines.
Palp of maxilla 1 bearing only 6-7 setae. Coxa 1
with sharper anterodistal corner than in female;
coxae 2-3 slightly wider than in female; coxa 4
slightly larger than in female; long setal formula of
coxae 1-4 = 7-8-8-3. Long posterior setae on article
2 of gnathopods 1-2 = 0-1, short anterior setae =
4 and 4 [long seta on gnathopod 2 possibly broken
off]; article 6 of gnathopods 1-2 slightly larger than
in female in relation to length and width of articles
3-5. Long posterior setae on article 2 of pereopods
1-2 = 1 and 4-5 (besides posteroventral pair), short
anterior setae = 4-3 and 4; article 4 with only one
small anterior subdistal seta on pereopod 1, none on
pereopod 2; article 6 with rows of one and one
posterior setae, rows of 5 and 5 spines on pereopod
1, 6 and 5 spines on pereopod 2. Article 2 of pereo-
pods 3-5 strongly to slightly narrower than in
female (see illustrations). Ridge on epimeron 2 ap-
parently absent, ventral setal row comprising 4 setae
tightly constricted; epimeron 3 with posteroventral
tooth obsolescent, larger midposterior teeth only 2
in number. Apicolateral spine on peduncle of uro-
pod 1 smaller than in female, ramal spines of uro-
pods 1-2 more numerous (see illustrations), medial
margin of peduncle on uropod 1 with one apical,
one mid spine and basal setules. Rami of uropod 3
masculine, inner extending almost to apex of article
1 on outer ramus, marginally setose, outer ram us
with 4 lateral spine groups. Telson longer than in
female, width-length ratio about 14:17, each lobe
with basal group of denticles, apices of lobes
sinuous, medial corner weakly produced.
VARIATIONS.?Female "n" with aberrant accessory
pair of deeply inserted and sheathed setae just
anteroventral to normal posterior pair of setae on
epimeron 2, left side only. Male "c" not fully de-
veloped; eyes enlarged; antenna 1 lacking calceoli
and large aesthetascs, primary flagellum 11-articu-
late, accessory flagellum 6 articulate, article 2 of an-
tenna 1 with 7 setae in main group; flagellum of
antenna 2 proliferate, .SO articulate, articles very
short, lacking calceoli; epimeron 2 with 5 setae;
telson it denticles absent: this specimen better de-
veloped than male "k" in terms of epimeron 2 and
articles of antenna I but otherwise immature.
ILLUSTRATIONS.?One setule on dorsal apex of
article 2 on antenna I of female hidden in illustra-
tion; right mandibular molar of female shown in
flattened view; on male antenna 2, fuzz on medial
face of article 4 concealed; left side of telson with
one spine missing.
HOLOTYPE.?NMV, female "a," 4.77 mm.
TYPE-LOCALITY.?CPBS 35S/3. 1 Mar 1965, West-
ern Port, Victoria, Australia, 14 m, sand.
VOUCHER MATERIAL.?Type-locality: female "n,"
5.04 mm (ilhis.); male "c," 4.93 mm; male "k," 4.82
mm (illus.).
MATERIAL.?CPBS, 28 samples from 11 stations
(63).
DISTRIBUTION.?Victoria, Western Port, 8-15 m,
sand.
HARPINIINAE
DIAGNOSIS.?Article 2 of antenna 1 especially
shortened; mandibular molar fully triturative or
reduced to a small hump with articulate spines;
palp of maxilla 1 uni- or biarticulate; setation on
maxilla 2 ordinary or reduced; gnathopod 2 sig-
nificantly enlarged or as small as gnathopod 1;
article 2 of pereopod 3 of narrow form; pereopod 5
usually with enlarged article 3.
DESCRIPTION.?Article 5 of antenna 2 usually re-
duced in size; epimeron 3 of nonrounded classifica-
tion; apices of peduncles on uropods 1-2 usually
not combed.
TYPE-GENUS.?Harpinia Boeck.
NUMBER 245 529
COMPOSITION.?Basuto, new genus; Coxophoxus
J. L. Barnard; Harpiniopsis Stephensen; Hetero-
phoxus Shoemaker; Proharpinia Schellenberg;
Pseudharpinia Schellenberg.
REMARKS.?All phoxocephalid genera with thin
article 2 on pereopod 3 are placed in this subfamily.
The Harpiniin Group of Genera
This group of genera is in poor classificatory
state. The seven genera do not express the known
diversity adequately, numerous species are so poorly
described that they cannot be determined gener-
ically, and a generic revision of the group cannot be
undertaken until the missing details on known
species are collected. Part of the problem involves
knowing the males adequately.
The Harpiniin group appears to have an evolu-
tionary flow commencing in the subantarctic
islands, antiboreal South America, and Antarctica
and flowing northward primarily by submergence
into the deep sea. During this evolutionary process
the eyes of primitive antiboreal littoral forms were
lost, even though one genus, Harpinia, reemerged
in the north Atlantic onto sublitoral shelves.
The primitive antiboreal members of the Har-
piniin group have continuous dorsal spination on
one or more rami of uropods 1-2 but this decays
rapidly to the north, although a few species north
of the subantarctic convergence show some vestiges
of this spination remaining on the inner ramus of
uropod 2.
Two of the dominantly antiboreal genera,
Heterophoxus and Pseudharpinia, have a strong
ensiform process on article 1 of antenna 2, whereas
the other dominantly subantarctic genus, Pro-
harpinia , lacks this process. In the type-species of
Proharpinia and Pseudharpinia, article 2 of pereo-
pod 5 is grossly toothed and although this charac-
teristic deteriorates even within Antarctica, it is
often seen in northern deep-sea harpiniins. The dis-
tinctive Harpinia, endemic to the North Atlantic,
retains the ensiform process typical of Pseud-
harpinia and often has species with grossly toothed
pereopod 5. Most species of the northern Harpini-
opsis (abyssal, mainly Pacific, scarce in northern At-
lantic) have lost the ensiform process but occasion-
ally bear the heavily toothed pereopod 5.
Whether or not the presence of eyes is a good
generic character remains a problem for extensive
study. The generic concept of and the lines of
descent from Proharpinia, Pseudharpinia, and
Heterophoxus into the world abyssal genus Har-
piniopsis are difficult matters. One may hypothesize
a polyphyletic descent from each of the three genera
to form various members of Harpiniopsis.
The minute details of mandibles especially, but
also of other attributes in Harpiniopsis, must be
studied before a better solution can be offered than
now extant. However, in order to constrain Har-
piniopsis more closely than has been done hereto-
fore, we remove many species from Harpiniopsis
and place them in Pseudharpinia. These species are
characterized by a strong ensiform process on an-
tenna 2 and continuously spinose inner rami of
uropod 2. In other respects these species may or may
not have outer rami of uropods 1-2 continuously
spinose and they vary considerably in the gross
dentition on article 2 of pereopod 5. Gross denti-
tion itself may have value in distinguishing genera.
We present a new key and new diagnoses for the
seven genera and condensed lists of species with
emphasis only on those having been described since
J. L. Barnard (1960).
Key to the Genera of Harpiniinae
1. Eyes present 2
Eyes absent 5
2. Antenna 2 strongly ensiform Heterophoxus
Antenna 2 not ensiform 3
S. Article 4 of pereopod 4 thick, dactyl of maxilliped very long, apical nail obsolescent
Basuto, new genus
Article 4 of pereopod 4 thin, dactyl of maxilliped short, stubby, apical nail highly elongate . . 4
4. Mandibular molar triturative, hands of gnathopods rectangular Coxophoxus
Mandibular molar simple, spinose, hands of gnathopods ovate Propharpinia
5. Flagellum of male antenna 2 short as in female, article 1 of primary flagellum of male
antenna 1 elongate and brushy Harpinia
530 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Flagellura of male antenna 2 highly elongate and proliferate, article 1 of primary flagellum
on male antenna 1 scarcely elongate, not brushy *
6. Some rami of uropods 1-2 with dorsal spines continuous to apex, article 2 of outer ramus of
uropod 3 short, antenna 2 usually ensiform P$eudharpinia
No rami on uropods 1-2 continuously spinose to apex, article 2 on outer ramus of uropod 3
elongate, antenna 2 scarcely or not ensiform HmrpbiioptU
Coxophoxus J. L. Barnard
Coxophoxus J. L. Barnard, 1966a:84.
DIAGNOSIS.?Eyes present. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae confined apically.
[PArticle 1 of antenna 2 not ensiform; article S with
2 setules;] facial spines on article 4 in one main row
or mostly apical; article 5 especially thin and short.
[PRight mandibular incisor with 3, 4 teeth]; molar
triturative, large to medium; palpar hump small.
Palp of maxilla 1 uniarticulate; inner plate naked.
Setation of maxilla 2 weak. Inner plate of maxil-
lipeds ordinary; apex of palp article 3 not pro-
tuberant, dactyl elongate, apical nail distinct, elon-
gate. Gnathopods weakly dissimilar, gnathopod 1
moderately and gnathopod 2 strongly enlarged;
article 5 of gnathopod 1 of ordinary length, short
and almost cryptic on gnathopod 2, without eusirid
attachment, palms oblique to transverse, hands of
gnathopods 1-2 broadened, poorly setose anteriorly.
Article 5 of pereopods 1-2 scarcely setose postero-
proximally. Article 2 of pereopod 3 of narrow form,
articles 4-5 of pereopods 3-4 narrow, article 2 of
pereopods 3-4 weakly setose posteriorly; pereopod
5 ordinary, article 2 naked ventrally, article 3 ordi-
nary, dactyl normal. Urosomite 1 generally naked;
urosomite 3 without dorsal hook or special process.
Peduncle of uropod 1 normally elongate, without
apicoventral spike, without special enlarged apico-
lateral-medial spine, [Ppeduncular apices of uropods
1-2 not combed]; inner ramus of uropod 1 with
marginal spines in one row, no rami continuously
spinose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, article 2 of outer ramus carry-
ing 2-3 short to long apical setae. Telson ordinary,
with only one apical spine on each lobe plus
setules, without special dorsal and lateral spines or
setae. Epimera 1-2 lacking long posterior setae,
without midfacial setae above ventral facial ridge;
epimeron 3 ordinary, almost of rounded classifica-
tion but bearing 4 or more long setae.
DESCRIPTION.?Rostrum fully developed. [Fuzz on
article 1 of antenna 1 in male unknown; calceoli on
male antennae 1-2 unknown], but flagellum of an-
tenna 2 scarcely proliferate or elongate. [Prebuccal
parts unknown. Right lacinia mobilis unknown;]
mandibular palp thin, article 1 short, article 2 with-
out outer setae, apex of article 3 transverse. Lower
lip lacking cones. Outer plate of maxilla 1 with 9
spines, [Pone spine especially thickened]. Inner
plates of maxilltped poorly armed, thick. Coxae 2-4
without special an termini sal humps. All posterior
spines on article 6 of pereopods 1-2 thin and seta-
like, [Pmidapical spine or seta present]. Article 2 of
pereopod 5 without facial setae. Peduncle of uropod
1 with dorsolateral spines widely spread, [Pmedial
spines confined apically; peduncle of uropod 2 with
only one medial spine confined apically]. Peduncle
of uropod 3 lacking extra subapical setae or spines.
[PTelson with ordinary pair of midlateral or dorsal
setules on each side.]
TYPE-SPECIES.?Coxophoxus hidalgo J. L. Bar-
nard, 1966a [originally designated].
COMPOSITION.?Phoxocephalus coxalis K. H.
Barnard, 1932.
Coxophoxus hidalgo J. L. Barnard
Coxophoxus hidalgo J. L. Barnard, 1966a:84, figs. 37, 38.
DISTRIBUTION.?Off southern California, 1675 m.
Coxophoxus coxalis (K. H. Barnard)
Phoxocephalus coxalis K. H. Barnard, 1932:97-99, fig. 48.
Coxophoxus coxalis.?J. L. Barnard, 1966a:84.
DISTRIBUTION.?South Georgia, 0-5 m.
Basuto, new genus
DIAGNOSIS.?Eyes present. Flagella of antenna 1-2
unreduced in female. Article 2 of antenna 1 espe-
cially shortened, ventral setae widely spread but
mostly apical. Article 1 of antenna 2 not ensiform;
article 3 with numerous setae and setules; facial
NUMBER 245 5S1
spines on article 4 in 2 rows set highly apicad; arti-
cle 5 especially short. Right mandibular incisor with
3 teeth; molar not triturative, small apparently
bearing 4 or more splayed, semiarticulate spines,
[Pusually not bearing fuzz]; palpar hump large. Palp
of maxilla 1 biarticulate; inner plate with 3 setae.
Setation of maxilla 2 ordinary. Inner plate of
maxillipeds ordinary; apex of palp article 3 not
protuberant, dactyl elongate, apical nail distinct,
medium. Gnathopods dissimilar, gnathopod 2 mod-
erately to strongly enlarged; article 5 of gnathopods
1-2 short, free on gnathopod 1, cryptic on gnatho-
pod 2, with weak eusirid attachment, palms oblique,
hands of both gnathopods elongate, broadened in
gnathopod 2, poorly setose anteriorly. Article 5 of
pcreopods 1-2 with posteroproximal setae. Article
2 of pereopod S of narrow form, articles 4-5 of
pcreopods 3-4 narrow to medium, article 2 of
pcreopods 3-4 not setose posteriorly; pereopod 5
ordinary; article 2 strongly setose ventrally; article
3 ordinary to weakly enlarged; dactyl normal. Epi-
mera 1-2 lacking long posterior setae, without mid-
facial setae above ventral facial ridge, epimeron 3
ordinary; urosomite 1 generally naked, urosomite 3
without dorsal hook or special process. Peduncle of
uropod 1 normally elongate, without apicoventral
spike, with special enlarged apicolateral spine,
[Ppeduncular apices of uropods 1-2 not combed],
inner ramus of uropod 1 with marginal spines in
row, some rami continuously spinose to apex, inner
ramus of uropod 2 especially shortened. Uropod 3
ordinary, elongate article 2 of outer ramus carrying
2 long apical setae. Telson ordinary, with only one
apical spine on each lobe plus setules.
DESCRIPTION.?Rostrum fully developed. [Fuzz
and calceoli on male antennae unknown. Prebuccal
parts unknown. Right lacinia mobilis unknown;]
mandibular palp medium to thick, article 1 short,
article 2 with outer setae, apex of article 3 oblique.
Lower lip bearing cones. Outer plate of maxilla 1
with 9 spines, one spine especially thickened. Inner
plates of maxilliped especially thick, ordinarily
setose. Coxae 2-4 without special anterodorsal
humps. All posterior spines on article 6 of pereopods
1-2 probably thin and stiff, [midapical spine or
seta unknown]. Article 2 of pereopod 5 without
facial setae. [PPedunde of uropod 1 with dorso-
lateral spines confined apically], medial spines con-
fined apically, peduncle of uropod 2 with 2 elon-
gate medial setae confined apically, peduncle of
uropod 3 lacking extra subapical setae or spines.
Telson with ordinary pair of midlateral or dorsal
setules on each side.
TYPE-SPECIES.?Pontharpinia stimpsoni Stebbing,
1908 (here selected; monotypic).
COMPOSITION.?Unique.
RELATIONSHIP.?Basuto resembles Mandibulo-
phoxus but differs in the mukisetose article 3 of
antenna 2, as in Pontharpinia, the highly narrowed
article 2 of pereopod 3, as in the harpiniin genera,
has a small inner ramus on uropod 2, two long
setae on the medial peduncle of uropod 2, a more
enlarged gnathopod 2 and a telson bearing only
one apical spine but with normal midlateral setule
pairs. This genus is named for a region in southern
Africa. Masculine.
Basuto stimpsoni (Stebbing), new combination
Pontharpinia stimpsoni Stebbing, 1908:75-78, pi. 11; 1910:
452.?Reid. 1951:222, fig. 22.
Pontharpinia Stimpsoni.?Chevreux, 1925:296.
Pontharpinia intermedia Schellenberg, 1925:138-140, fig. 11.
?Reid, 1951:223.
Mandibulophoxus stimpsoni.?J. L. Barnard, 1957:436-438,
fig. 4; 1960:359.?Griffiths, 1974a:250; 1974b:321.
DISTRIBUTION.?Africa, from Senegal southward
around South Africa to the Natal Coast, 5-300 m.
Proharpbua Schellenberg
Proharpinia Schellenberg, 1931:80.?J. L. Barnard, 1960:311.
DIAGNOSIS OF TYPE-SPECIES.?Eyes present.
Flagella of antennae 1-2 unreduced in female.
Article 2 of antenna 1 especially shortened, ventral
setae widely spread or confined apically. Article 1
of antenna 2 weakly ensiform; article 3 with
numerous setae and setules; facial spines on article
4 in one main row; article 5 especially short.
[?Right mandibular incisor with 3 teeth]; molar
not triturative, ordinary, small, pillow-shaped, bear-
ing 3 clumped spines [Pwith common base, usually
not bearing fuzz]; palpar hump small. Palp of
maxilla 1 biarticulate; inner plate naked. Setation of
maxilla 2 ordinary. Inner plate of maxillipeds
ordinary; apex of palp article 3 not protuberant,
dactyl short, stubby, apical nail distinct, elongate.
Gnathopods ordinary, small, similar; article 5 of
gnathopods 1-2 short, free, without eusirid attach-
532 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ment, palms oblique, hands of gnathopods 1-2
ordinary, ovatorectangular to elongate, poorly
setose anteriorly. Article 5 of pereopods 1-2 setose
posteroproximally. Article 2 of pereopod 3 of nar-
row form, articles 4-5 of pereopods 3-4 narrow to
medium, article 2 of pereopods 3-4 not setose
posteriorly; pereopod 5 ordinary, article 2 strongly
toothed ventrally, article 3 ordinary, dactyl normal.
Epimera 1-2 lacking long posterior setae, without
midfacial setae above ventral facial ridge; epime-
ron 3 ordinary. Urosomite 1 generally naked; uro-
somite 3 without dorsal hook or special process.
Peduncle of uropod 1 normally elongate, without
apicoventral spike, without special enlarged apico-
lateral-medial spine, [Ppeduncular apices of uro-
pods 1-2 not combed], inner ramus of uropod 1
with marginal spines in one row, some rami con-
tinuously spinose to apex, inner ramus of uropod
2 ordinary. Uropod 3 ordinary, elongate article 2
of outer ramus [Pcarrying vestigial apical setae].
Telson ordinary, with only one apical spine or seta
on each lobe, without special dorsal and lateral
spines or setae.
DESCRIPTION.?Rostrum fully developed, head
with anteroventral tooth. [Fuzz on article 1 of an-
tenna 1 in male unknown; calceoli on male anten-
nae 1-2 unknown. ?Prebuccal parts ordinary. Right
lacinia mobilis unknown;] mandibular palp thin,
article 1 short, article 2 without outer setae, apex
of article 3 oblique. [PLower lip bearing cones.]
Outer plate of maxilla 1 with 9 spines, [?no spine
especially thickened]. Inner plates of maxilliped
especially thick, ordinarily setose. Coxae 2-4 with-
out special anterodorsal humps. [?A11 posterior
spines on article 6 of pereopods 1-2 thin and seta-
like, midapical spine or seta present.] Article 2 of
pereopod 5 without facial setae. Peduncle of uro-
pod 1 with dorsolateral spines widely spread, me-
dial spines widely spread, [Ppedunde of uropod 2
with only one medial spine or setule confined api-
cally]. Peduncle of uropod 3 lacking extra sub-
apical setae or spines. [PTelson with midlateral or
dorsal setules reduced to one member on each side.]
TYPE-SPECIES.?Proharpinia antipoda Schellen-
berg, 1931 (monotypic).
COMPOSITION.?Proharpinia hurleyi J. L. Bar-
nard, 1958; Heterophoxus stephenseni Schellen-
berg, 1931; Proharpinia tropicana J. L. Barnard,
1960 (See J. L. Barnard (1960) for references and
distribution).
REMARKS. None of the three species that have
been assigned to this genus agree with the type-
species in the continuous dorsal spination on the
inner ramus of uropod 2 and none oi them has
a grossly toothed article 2 on pereopod 5. Two
of the species, P. stephemerii and P. tropicana,
have a poorly toothed epimeron 3. If these 3 spe-
cies lacked eyes they would fall to Harpiniopsis.
Heterophoxus Shoemaker
Heterophoxus Shoemaker, 1925:22 ?J L. Barnard. 1960:318.
DIAGNOSIS.?Eyes present. Flagella of antenna
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae widely spread
towards apical end. Article 1 of antenna 2 strongly
ensiform; article 3 with numerous setules: facial
spines on article I in one main row; article 5 espe-
cially thin and short. Right mandibular incisor
with 4+ teeth; molar not triturative, ordinary,
small, pillow-shaped, bearing 3 long clumped
spines, not bearing fuzz; palpar hump medium.
Palp of maxilla 1 biarticulate; inner plate with 2
setae. Setation of maxilla 2 ordinary to weak. Inner
plate of maxillipeds partly fused together, apex of
palp article 3 not protuberant, dactyl short, stubby,
apical nail distinct, elongate. Gnathopods small,
similar; article 5 very short, almost cryptic, without
eusirid attachment, palms oblique, hands ovatorec-
tangular, elongate, poorly setose anteriorly. Article
5 of pereopods 1-2 setose proximoposteriorly. Arti-
cle 2 of pereopod 3 of narrow form, articles 4-5
of pereopods 3-4 narrow, article 2 of pereopods
3 or 4 setose posteriorly; pereopod 5 ordinary, arti-
cle 2 strongly setose and toothed ventrally, article
3 slightly enlarged, dactyl normal. Epimera 1-2
lacking numerous long posterior setae, without
midfacial setae above ventral facial ridge; epimeron
3 ordinary. Urosomite 1 generally naked; uroso-
mite 3 without dorsal hook or special process. Pe-
duncle of uropod 1 normally elongate, without
apicoventral spike, without special enlarged apico-
lateral-medial spine, only peduncular apex of uro-
pod 2 combed, inner ramus of uropod 1 with mar-
ginal spines in one row, either none or some rami
continuously spinose to apex, inner ramus of uro-
pod 2 ordinary. Uropod 3 ordinary, small article 2
of outer ramus carrying 2 long apical setae. Telson
ordinary, with only one apical spine or seta on each
lobe, often with special lateral spine on each side.
NUMBER 245 533
DESCRIPTION.?Rostrum fully developed. Fuzz on
article 1 of antenna 1 in male present, calceoli
present on male primary flagellum of antenna 1,
and on article 5 and flagellum of male antenna 2.
Prebuccal parts extended forward, massive, strongly
distinct, epistome dominant. Right lacinia mobilis
bifid, flabellate; mandibular palp medium to thin,
article 1 short, article 2 without outer setae, apex
of article 3 oblique. Lower lip bearing cones. Outer
plate of maxilla 1 with 11 spines, one spine espe-
cially thickened. Inner plates of maxilliped poorly
armed, thick. Coxae 2-4 without special antero-
dorsal humps. All posterior spines on article 6 of
pereopods 1-2 thin and seta-like, midapical spine
or seta absent (but apicolateral spine very short
and pointed inward). Article 2 of pereopod 5 with-
out facial setae. Peduncle of uropod 1 with dorso-
lateral spines confined apically, medial spines
widely spread, peduncle of uropod 2 with about 2
medial spines widely spread. Peduncle of uropod 3
lacking extra subapical setae or spines. Telson with
ordinary pair of midlateral or dorsal setules on
each side.
TYPE-SPECIES.?Harpinia oculata Holmes, 1908
(= Heterophoxus pennatus Shoemaker, 1925)
(monotypic); now Heterophoxus oculatus (Holmes,
1908).
COMPOSITION.?Harpinia ophthalmica Schellen-
berg, 1925; Heterophoxus trichosus K. H. Barnard,
1932; Heterophoxus videns K. H. Barnard, 1930
(see J. L. Barnard (1960) for references and distri-
bution).
REMARKS.?The diagnostic details have been con-
firmed on specimens of the type-species deposited
in the Smithsonian Institution.
Pseudharpinia Schellenberg
Pseudharpinia Schellenberg, 1931:81-82.?J. L. Barnard, 1960:
342.
DIAGNOSIS, (based on type-species).?Eyes absent.
Flagella of antennae 1-2 unreduced in female. Arti-
cle 2 of antenna 1 especially shortened, ventral
setae widely spread. Article 1 of antenna 2 strongly
ensiform; article 3 with numerous setae and set-
ules; facial spines on article 4 in one main row
or 2 poorly denned rows; article 5 especially thin
and short. [PRight mandibular incisor with 4 teeth];
molar not triturative, medium, pillow-shaped,
bearing 4 or more splayed, semiarticulate spines,
[Pusually not bearing fuzz]; palpar hump small.
Palp of maxilla 1 biarticulate; inner plate with 2
setae. Setation of maxilla 2 weak. Inner plate of
maxillipeds ordinary, apex of palp article 3 not
protuberant, dactyl weakly elongate, somewhat
stubby, apical nail distinct, elongate. Gnathopods
small, similar, but gnathopod 2 weakly enlarged;
article 5 of both gnathopods very short, free, with
weak eusirid attachment, palms oblique, hands
ordinary, ovatorectangular, weakly elongate, poorly
setose anteriorly. Article 5 of pereopods 1-2 setose
posteroproximally. Article 2 of pereopod 3 of nar-
row form, articles 4-5 of pereopods 3-4 narrow,
article 2 of pereopods 3-4 setose posteriorly; pereo-
pod 5 ordinary, article 2 strongly setose and toothed
ventrally, article 3 enlarged, dactyl normal. Epi-
mera 1-2 lacking long posterior setae, without mid-
facial setae above ventral facial ridge; epimeron 3
ordinary. Urosomite 1 generally naked; urosomite
3 without dorsal hook or special process. Peduncle
of uropod 1 normally elongate, without apicoven-
tral spike, without special enlarged apicolateral-
medial spine, [Ppeduncular apices of uropods 1-2
not combed], inner ramus of uropod 1 with mar-
ginal spines in 2 rows, some rami continuously spi-
nose to apex, inner ramus of uropod 2 ordinary.
Uropod 3 ordinary, small article 2 of outer ramus
carrying 2 long apical setae. Telson with 2-4 apical
spines or setae on each lobe, often with special
dorsal and lateral setae.
DESCRIPTION.?Rostrum fully developed. [Fuzz on
article I of antenna 1 in male unknown, calceoli
on male antennae 1-2 unknown. PPrebuccal parts
ordinary. Right lacinia mobilis unknown;] man-
dibular palp thin, article 1 short, article 2 without
outer setae, apex of article 3 oblique. Lower lip
bearing cones. Outer plate of maxilla 1 with 9
spines, one spine especially thickened. Inner plates
of maxilliped thick, ordinarily setose. Coxae 2-4
without special anterodorsal humps. [PA11 posterior
spines on article 6 of pereopods 1-2 thin and seta-
like, midapical spine or seta present]. Article 2 of
pereopod 5 without facial setae. Peduncle of uro-
pod 1 with dorsolateral spines widely spread, me-
dial spines confined apically; peduncle of uropod
2 with only one medial spine or setule confined
apically. Peduncle of uropod 3 lacking extra sub-
apical setae or spines. Telson with ordinary pair
of midlateral of dorsal setules on each side, highly
apicad.
534 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TYPE-SPECIES.?Pseudharpinia den tat a Schellen-
berg, 1931 (monotypic).
COMPOSITION.?Harpinia abyssalis Pirlot, 1932;
Harpinia ayutlanta J. L. Barnard, 1964b (Pacific
Panama, 1609-1746 m, ensiform process poorly de-
veloped); fHarpinia birjulini Gurjanova, 1953 (but
uropod 2 unknown); Harpinia brevirostris Chev-
reux, 1920, 1927; Harpinia cariniceps K. H. Barn-
ard, 1932 (but uropod 2 unknown); Harpinia cinca
J. L. Barnard, 1961, 1962 (Southwest Africa, 990-
4680 m); Harpinia excavata Chevreux, 1887, 1900
(= Harpiniopsis sanpedroensis J. L. Barnard, 1960);
Harpinia latipes Norman, 1900; Harpinia obtusi-
jrons Stebbing, 1888; Harpinia vallini Dahl, 1954
(Ross Sea, 550 m) (for references and distribution
consult J. L. Barnard (1960) except for those de-
scribed or allocated subsequently).
Rejected species: Harpinia palabria J. L. Barn-
ard, 1961 (Tasman Sea, 610 m), ensiform process
unknown but uropod 1 with spike, epimera 1-2
with posterior setae, to be described in a new
genus (J. L. Barnard, in prep.)
REMARKS.?Pseudharpinia excavata bears wide
medial spination on the peduncles of uropods 1-2;
the following species have poorly ensiform antenna
2: Harpinia ayutlanta, H. cinca; the following spe-
cies have elongate dactyl on the maxilliped: H.
abyssalis, H. obtusifrons, and H. spaercki (the lat-
ter assigned to Harpiniopsis).
Pseudharpinia dentata Schellenberg
Pseudharpinia dentata Schellenberg, 1931:82-83, fig. 42 ?
J. L. Barnard, 1960:342, pi. 75.
DISTRIBUTION.?Lagotowia.
Harpiniopsis Stephensen
Harpiniopsis Stephensen, 1925:171.?J. L. Barnard, 1960:325.
DIAGNOSIS.?Eyes absent. Flagella of antennae
1-2 unreduced in female. Article 2 of antenna 1
especially shortened, ventral setae confined apically.
Article 1 of antenna 2 not or weakly ensiform;
article 3 with numerous setae and setules; facial
spines on article 4 in one main row; article 5 ordi-
nary in size to especially short. Right mandibular
incisor with 3-4 teeth; molar not triturative, small,
pillow-shaped, bearing 2 or more splayed, semi-
articulate spines, usually not bearing fuzz; palpar
hump small to medium. Palp of maxilla 1 biartic-
ulate. Inner plate with 2 setae. Setation of maxilla
2 weak. Inner plate of maxillipeds ordinary, apex
of palp article 3 not protuberant, dactyl short,
stubby, apical nail distinct, elongate. Gnaihopods
ordinary, small, similar or gnathopod 2 weakly en-
larged; article 5 of both gnathopods very short,
almost cryptic on gnathopod 2, without eusirid
attachment, palms oblique, hands ordinary, ovato-
rectangular, elongate, poorly setose anteriorly. Arti-
cle 5 of pereopods 1-2 setose posteroproximally.
Article 2 of pereopod 3 of narrow form, articles
4-5 of pereopods 3-4 narrow, article 2 of pereopods
3-4 not setose posteriorly; pereopod 5 ordinary,
article 2 naked or weakly to strongly toothed ven-
trally, article 3 enlarged, dactyl normal. Epimera
1-2 lacking long posterior setae, without midfacial
setae above ventral facial ridge; epimeron 3 ordi-
nary or of rounded classification bearing fewer
than 3 setae. Urosomite 1 generally naked; uroso-
mite 3 without dorsal hook or special process. Pe-
duncle of uropod 1 normally elongate, without
apicoventral spike, without special enlarged apico-
lateral-medial spine, [?peduncular apices of uro-
pods 1-2 not combed], inner ramus of uropod 1
with marginal spines in one row, no rami contin-
uously spinose to apex, inner ramus of uropod 2
ordinary. Uropod 3 ordinary, elongate article 2 of
outer ramus carrying 1-2 short to long to vestigial
apical setae. Telson with 1-4 apical setae on each
lobe plus setules, occasionally with special dorsal
and lateral setae.
DESCRIPTION.?Rostrum fully developed, head
often with anteroventral tooth. [Fuzz on article 1 of
antenna 1 in male unknown; calceoli on male pri-
mary flagellum of antenna 1 unknown.] Calceoli
on article 5 of male antenna 2 present; flagellum
in male elongate, [?flagellum in male with calceoli].
Prebuccal parts ordinary to strongly extended for-
ward, occasionally massive, strongly distinct or
poorly separated from each other, dominance of
either epistome or upper lip variable. Right lacinia
mobilis bifid or simple, flabellate or thin; man-
dibular palp thin, article 1 short, article 2 without
outer setae, apex of article 3 oblique. Lower lip
bearing cones. Outer plate of maxilla 1 with 11
spines, one spine especially thickened. Inner plates
of maxilliped poorly armed, thick. Coxae 2-4 with-
out special anterodorsal humps. All posterior
spines on article 6 of pereopods 1-2 thin and seta-
NUMBER 245 535
like, [Pmidapical spine or seta present]. Article 2
of pereopod 5 without facial setae. Peduncle of
uropod 1 with dorsolateral spines confined apically,
medial spines confined apically or widely spread;
peduncle of uropod 2 with only one medial spine
or setule confined apically. Peduncle of uropod 3
lacking extra subapical setae or spines. Telson with
pair of midlateral or dorsal setules on each side,
highly apicad.
TYPE-SPECIES.?Harpiniopsis similis Stephensen,
1925 (monotypic).
COMPOSITION.?mHarpinia amundseni Gurjanova,
1946; ? Harpinia australis J. L. Barnard, 1961
(Great Australian Bight, 1320-1340 ra); ?Harpini-
opsis capensis (J. L. Barnard, 1962) (as Harpinia
laevis capensis, new subspecies, South Africa, 4893-
4961 m); ?Harpiniopsis emeryi J. L. Barnard, 1960;
Harpiniopsis epistomata J. L. Barnard, 1960; Har-
piniopsis fulgens J. L. Barnard, 1960; Harpiniopsis
galera J. L. Barnard, 1960; ?Harpinia gurjanovae
Bulycheva, 1936; *Harpinia kobjakovae Bulycheva,
1936; ?Harpinia miharaensis Nagata, 1960 (Japan,
shallow water); *Harpinia moiseevi Gurjanova,
1953; ?Harpinia nadania J. L. Barnard, 1961 (Tas-
man Sea, 610 m); * Harpiniopsis naiadis J. L. Barn-
ard, 1960; *Harpinia orientalis Bulycheva, 1936;
?Harpinia pacifica Bulycheva, 1936; Harpiniopsis
percellaris j . L. Barnard, 1971 (Oregon, 600-2600
m); *Harpiniopsis petitions J. L. Barnard, 1966a
(California, 1265 m); Harpiniopsis profundis J. L.
Barnard, 1960; *Harpinia salebrosa Gurjanova,
1936; mHarpinia schurini Bulycheva, 1936; *Har-
pinia spaercki Dahl, 1959 (Banda Trench, 6580-
7270 m); *Harpinia tarasovi Bulycheva, 1936;
?Harpiniopsis triplex J. L. Barnard, 1971 (Oregon,
2000-2800 m); *Harpinia wandichia J. L. Barnard,
1962 (South Sandwich Trench, 2747 m); ?Harpini-
opsis sp. D, J. L. Barnard, 1960. (See J. L. Barnard
(1960) for references and distribution, many of
these species placed with Harpinia at that time;
others, with literature and distribution subse-
quently described, as noted; asterisks denote species
of which the males remain undiscovered, so that
firm generic identification is impossible.)
REMARKS.?Harpinia laevis capensis J. L. Bar-
nard, 1962, is elevated to full specific level because
the species cannot be Harpinia laevis from the
North Atlantic Ocean owing to the absence of an
ensiform process on antenna 2. Harpiniopsis emeryi
bears a weak ensiform process, thereby demonstrat-
ing intergradation with ancestral antiboreal taxa.
See Pseudharpinia for species transferred to that
genus.
The species of this genus are poorly known and
may be divisible into additional genera. The diag-
nosis presented above is conceptually based on
knowledge of several species but not fully on any
one species, including the type-species.
Harpinia Boeck
Harpinia Boeck. 1876:218.?San, 1895:150?Stebbing, 1906:
140.?J. L. Barnard, 1960:344.
DIAGNOSIS.?Eyes absent. Flagella of antennae
1-2 unreduced in female, but male antenna 2 with
short flagellum. Article 2 of antenna 1 especially
shortened, ventral setae confined apically. Article 1
of antenna 2 strongly ensiform; article 3 with 2
setules; facial spines on article 4 in one main row;
article 5 especially thin and short. Right tnan-
dibular incisor with 4 teeth; molar not tritura-
tive, small, pillow-shaped, bearing 3 or more
splayed, semiarticulate spines, usually not bearing
fuzz; pal par hump small. Palp of maxilla 1 bi-
articulate; inner plate with 2 setae. Setation of
maxilla 2 weak. Inner plate of maxillipeds ordi-
nary, apex of palp article 3 weakly protuberant,
dactyl weakly elongate to short and stubby, apical
nail distinct, elongate. Gnathopods weakly dissimi-
lar, gnathopod 2 weakly enlarged; article 5 of both
gnathopods very short, free, with weak eusirid
attachment, palms oblique, hands ordinary, ovato-
rectangular, poorly setose anteriorly. Article 5 of
pereopods 1-2 setose posteroproximally. Article 2
of pereopod 3 of narrow form, articles 4-5 of pe-
reopods 3-4 narrow, article 2 of pereopods 3-4 not
setose posteriorly; pereopod 5 ordinary, article 2
naked or strongly toothed ventrally, article 3 en-
larged, dactyl normal. Epimera 1-2 lacking long
posterior setae, without midfacial setae above ven-
tral facial ridge; epimeron 3 ordinary or of rounded
classification and bearing 3 or fewer long setae.
Urosomite 1 generally naked; urosomite 3 without
dorsal hook or special process. Peduncle of uropod
1 normally elongate, without apicoventral spike,
without special enlarged apicolateral-medial spine,
peduncular apices of uropods 1-2 not combed;
inner ramus of uropod 1 with marginal spines in
one row, no rami continuously spinose to apex,
inner ramus of uropod 2 ordinary. Uropod 3 ordi-
536 SMITHSONIAN CONTRIBUTION'S TO ZOOLOGY
nary, elongate to short article 2 of outer ramus
carrying 1-2 long to vestigial apical setae. Telson
ordinary, with 1-4 apical spines or setae on each
lobe plus setules, rarely with special dorsal and
lateral setae.
DESCRIPTION.?Rostrum fully developed, head
often with anteroventral tooth. Fuzz on article 1
of antenna 1 in male present; calceoli on male pri-
mary flagellum of antenna 1 absent, article 1 of
primary flagellum elongate and fuzzy in male. Cal-
ceoli on article 5 of male antenna 2 present; flagel-
lum in male without calceoli. Prebuccal parts ordi-
nary, strongly distinct, both epistome and upper
lip prominent. Right lacinia mobilis bifid, flabel-
late or thin, mandibular palp medium to thin,
article 1 short, article 2 without outer setae, apex
of article 3 oblique. [PLower lip lacking cones.]
Outer plate of maxilla 1 with 7-9 spines, one spine
occasionally thickened. Inner plates of maxilliped
poorly armed, thick. Coxae 2-4 without special
anterodorsal humps. All posterior spines on article
6 of pereopods 1-2 thin and seta-like, [Pmidapical
spine or seta present]. Article 2 of pereopod 5 with-
out facial setae. Peduncle of uropod 1 with dorso-
lateral spines widely spread, [Pmedial spines con-
fined apically; peduncle of uropod 2 with only one
medial spine or setule confined apically]. Peduncle
of uropod 3 lacking extra subapical setae or spines.
Telson with ordinary pair of midlateral or dorsal
setules on each side. Mouth parts occasionally de-
generate in male.
TYPE-SPECIES.?Phoxus plumosus Kr0yer, 1842
(originally designated); now Harpinia plumosa
(Kr0yer, 1842).
COMPOSITION.?Harpinia abyssi Sars, 1885: Har-
pinia antennaria Meinert, 1893 (= H.ncglccta Sars,
1895); Harpinia bidentata Stephensen, 1925; Har-
pinia cabolemis Shoemaker. 1930; Harpinia crenu-
lata Boeck, 1871; fHarpinia crenuloides Stephen-
sen, 1925 (male unknown but antenna 2 with ensi-
form process); Harpinia curtipes Stephensen, 1925;
Harpinia dellavallei Chevreux, 1911; Harpinia lae-
vis Sars, 1895; Harpinia mucronata San, 1885;
Harpinia pectinata Sars, 1895; Harpinia propinqua
Sars, 1895, Bousfield, 1973; Harpinia serrata Sars,
1882; Harpinia truncata Sars. 1895 (see J. L. Barn-
ard, 1960, for references and distribution).
REMARKS.?This genus is confined to the North
Atlantic region as far as is known. However, the
males of many North Pacific species have not been
described and there is the possibility that species
now assigned to Harpiniopsis may belong with
Harpinia. The male of Harpinia is very unusual
in the shortness of the flagellum of antenna 2 that
apparently lacks calceoli (but calceoli are present
on the peduncle), the largeness of the brushes on
the antennae and the elongate and brushy article
1 on the primary flagellum of antenna 1.
Appendix
Australian Samples and Localities
AM. The Australian Museum, New South Wales, Australia,
samples and localities:
P.34S8. Port Jackson, New South Wales.
G.54IS. Port Jackson, New South Wales.
G.5406. Bondi Bay, Sydney, New South Wales.
P.18l2.f>. Antechamber Bay, Kangaroo Island, South Austra-
lia, with light in net, CSIRO Fisheries, 12 Dec 1939.
P.18I75. Off Tatmanian coast, dredged, 100 fathoms, C.
Hedley.
P.18I87. S of Cape Everard, Victoria. 58?12.5'S, 149?05.5'E.
152 in. per L. R. Thomas, 20 Jun 1962.
P. 18210. S of Cape Everard. Victoria, 37?55'S, I49?E, 77 m,
CSIRO Fisheries, per L. R. Thomas, 20 Jun 1962.
P.I82I4. Long Reef. Collaroy, New South Wales, P. Hutch-
ingt. 16 Nov 1970.
P. 18215. Off Sydney. New South Wales. 33e58.4'S, 151?
29% 150 m. Gear OPG. CSIRO Fisheries, per L. R.
1 hoinas 18 Jun 1962.
P.182I9. Same as P.18215.
P.18222. Same as P.18215.
P.18228. 70 mites S of Lakes Entrance, Victoria, 148?2450T.
S9?S, sand, 52 fathoms, "Esso-Gipps" Sta 20, per C.
Phipps. May 1969.
P. 18229. Off island in middle of Balmoral Head, Port
Jackson, New South Wales, 20 ft [6.1 m], mud bottom,
N. Coleman, 27 Jun 1970.
P.18242. Same as P.18215.
P.18243. Same as P.18215.
P. 18248. Same as P.18215.
P.18253. Same as P.18215.
P. 18269. SW of Cape Everard, Victoria, 38?15'S, 148?12'E,
300 m, Gear OPG, CSIRO Fisheries, per L. R. Thomas,
20 Jun 1962.
P. 18270. Same as P.18269.
P.18276. S of Eucla. Western Australia, Great Australian
Bight, 33?05'S, 128?40'E, 83 m, CSIRO Fisheries, per
L. R. Thomas, 5 Jul 1962.
P. 18285. Off Manly Beach, Sydney, New South Wales,
trawled 18-20 ft [53-6.1 m], bottom dean sand, P.
Hutchings, 30 Jun 1971.
P.18312. E of Hood Point, Western Australia, 34?32'S, 121?
31'E, 145 m. CSIRO Fisheries, 9 Jul 1962.
CPBS. Victoria Division of Fishieries and Wildlife, Crib
Point Benthic Survey, in Western Port. Samples were taken
at 52 stations (Figure 4) between 19 Feb and 13 Apr 1965,
with periodic samplings at stations 300, 31N, 31S, 32N, 32S
until 1970, and occasional resampling at some other sta-
tions. Phoxocephalids were found at every station and were
present in 293 of a total of 326 samples. A small prelimi-
nary survey of the Crib Point area (Figure 4. inset), was
made between 9 Jul and 13 Oct 1964; 43 samples were
examined, all containing phoxocephalids.
EBS. New South Wales State Fisheries, Estuarine Benthic
Survey, (Figure 1, insets 1 and 2), samples and localities:
20. Jervis Bay, Murray's Beach, 6 m, Zostera, 25 Apr 1972.
22. Same as 20.
27. Jervis Bay, Murray's Beach, 6 m, sand, 25 Apr 1972.
28. Same as 27.
30. Same as 27.
31. Jervis Bay, Murray's Beach, 6 m, Halophila, 25 Apr
1972.
32. Same as 31. <
72. Botany Bay, entrance to Woolaware Bay, mud, shell-
grit, 2.5 m, 14 Sep 1972.
103. Georges River, Woronora, 3 m, sand, 25 Oct 1972.
117. Georges River, Panama, 3 m, sand and mud, 25 Oct
1972.
138. Botany Bay, Towra Beach, 0.9 m, Zostera, shell-grit,
30 Nov 1972.
148. Botany Bay, 500 yds (457 m) off Towra Point, 3 m,
sand, shell-grit, 12 Jan 1973.
149. Same as 148.
196. Botany Bay, off Towra Beach, 3 m, sand, 11 Apr 1973.
199. Same as 196.
200. Same as 196.
304. Botany Bay. Towra Point, 0.9 m, Zostera, 17 Apr 1973.
311. Same as 304.
313. Same as 304.
362. Lake Macquarie, Salt's Bay, 0.3 m, coarse sand, 10 Jul
1973.
363. Same as 362.
JLB AUS. Samples collected by J. L. Barnard in Western
Australia with localities:
3. Sugarloaf Rock, Cape Naturaliste, Western Australia,
inter tidal wash of common seaweeds, 1 Sep 1968.
11. Middleton Beach, Albany, Western Australia, inter-
tidal wash of algae and rocks, 30 Sep 1968.
12. Same as JLB AUS 11. coralline algae.
IS. Middleton Beach, Albany, Western Australia, intertidal
wash of sandy rocks, coralline algae, 30 Sep 1968.
Lake King. Specimens from hand samples collected at Lake
King, Gippsland, by J. Kudenov, Marine Pollution Studies
Group, Victoria Division of Fisheries and Wildlife.
Macreadie. Plankton sample 710131, collected by V. Macrea-
die, Monash University, Crib Point in Western Port, 8 Aug
1972.
537
538 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
MMD. Samples collected by Margaret M. Drummond as fol-
lows: Merimbula, New South Wales, intertidal shallows,
10 and 16 Feb 1972.
NMV. National Museum of Victoria, Melbourne, Victoria,
Australia: all holotype and voucher materials from the
surveys of Western Port and Port Phillip Bay are deposited
in this museum; in addition a sample with a phoxo-
cephalid from the Port Phillip Survey, 1957-63 is included
in this study.
PPBES. Victoria Division of Fisheries and Wildlife, Benthic
Survey in the Port Phillip Bay Environmental Study. Sam-
pling was carried out at 112 stations between 17 Oct 1969
and 2 Nov 1972 (Figure 2). Of a total of 452 samples, 314
contained phoxocephalids but they were not taken at sta-
tions 918, 933, 935, 940, 945, 947, 952, 954. 956, 958, 962.
RHM. Plankton samples collected by Ralph H. Miller, Uni-
versity of Melbourne, in the Crib Point area of Western
Port, 19 Jan 1968, 25 Mar 1968. 22 Oct 1971.
SBS. The Australian Museum, Shelf Benthic Survey, off
Malabar, New South Wales, (Figure 1, inset 2), samples
and localities:
A1S1. 31 m, coarse gravel, sand, shell fragments, 12 May
1972.
A2S3. 48 m, medium fine sand, 15 May 1972.
A2S5. Same as A2S3.
C1S5. 29 m, shelly gravel, high organic content, 17 May
1972.
C2S2. 69 m, sandy mud, medium fine sand, 17 May 1972.
C4S4. 69 m, medium fine sand, 17 May 1972.
C6S2. 75 m, sandy gravel, 22 May 1972.
C6S3. Same as C6S2.
C7(- Cx)Sl. 92 m, sandy gravel, 22 May 1972.
C7(? Cx)S4. Same as C7S1.
D1S1. 31 m, medium sand and shell fragments, 17 May
1972.
D1S2. Same as D1S1.
D1S3. Same as D1SI.
D1S4. Same as D1S1.
D2S1. 49 m, medium sand and shell fragments, 17 May
1972.
D2S2. Same as D2S1.
D2S3. Same as D2S1.
D2S4. Same as D2S1.
D2S5. Same as D2S1.
D3S3. 71 m, medium fine sand, 19 May 1972.
E2S3. 53 m, medium fine sand, 19 May 1972.
E2S4. Same as E2S3.
E4S1.66 m, gravel with medium sand, 19 May 1972.
E4S3. Same as E4S1.
3. 1.6 km E of Malabar outlet, Sydney. 150?16'E. 33?58'42"
S, 66 m, 23 May 1973.
4. 2 km E of Long Bay. Sydney. 15O?I7'E. 33?58'5O-S. 66
m. 23 May 1973.
5. 23 km E of Malabar. I5O?16'48"E. SS'bWTS. 66 m.
23 May 1973.
Little Bay. Malabar, 16.7 ro. gravel and sand, 19 Jun 1972.
Long Reef, Collaroy. 43 m. 27 Apr 1972.
Off North Head. Sydney. 33 m. 23 May 1972.
Station P. 29 m, gravel and sand. 27 Sep 1972.
Dredge 20. no data.
Shepherd 15. Sample collected by S. A. Shepherd. Pearson
Islands, South Australia, 8 in. 2/11 m*. horizontal face in
algae, Jan 1969.
Slack-Smith I. Sample collected by S. Slack-Smith. Albany,
Cheyne Beach, Western Australia, on weedy rocks, inter-
tidal. 4 Dec 1968.
L'SNM. Former United Stales National Museum (specimens
now in National Museum of Natural History, Smithsonian
Institution. Washington. D.C., U.S.A.); duplicates of spe-
imens from Western Port and Port Phillip Bay surveys
deposited in NMNH.
WAM. Western Australian Museum. Perth. Western Aus-
tralia, samples and localities:
9561. Off Garden Island, Western Australia, night tow, Feb
1920.
414-73. Burrow Island, Western Australia, night taw, 2 Sep
1954.
415-73. Trawl station off water run, 35 m, sand, 3 Feb
1943.
416-73. Trawl station 6, off Jibbon Point, New South
Wales, 24 Jul 1943.
417-73. Eden, New South Wales, 30 m. coarse sand. 8 Oct
1943.
419-73. Antechamber Bay, Kangaroo Island, South Austra-
lia, light in net at night.
WPBES. Victoria Ministry for Conservation, Division of
Fisheries and Wildlife, Westernport Bay Environmental
Study. Benthic samplings were made at 41 stations (Figure
3) between 20 Nov 1973 and 25 Jan 1974 in Western Port;
60, of a total of 123 samples, contained phoxocephalids but
they were not found at stations 1702, 1703, 1708, 1710,
1711, 1712, 1713, 1719, 1720, 1723, 1737, 1738. Material from
9 stations in the Western Entrance sampled in 1975, has
been examined, but full Western Entrance data are not
included here, though some specimens have been used.
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Index
(Names of \alid species and genera of Phoxocephalidae and also page numbers for major treat-
ment are in italics; the absence of italics for the specific or generic name (or both) in a binominal
Uxon of Phoxocephalidae indicates a classification change that is accomplished, propised, or
contemplated.)
abyssalis, Harpinia, 534
Pseudharpinia, 534
abyui, Harpinia, 536
abronius, Paraphoxus. 145
Amphibolus antarctica. SO
amundseni, Harpinia, 535
Hatpiniopiii. 5)5
Anisogammarut, 22
antarctica. Amphibolui, 30
antennaria, Harpinia. 5)6
ttntipoda, Proharpinia, 5)2
apart, Birubius, 196. 198. 200. 201. 228, 2S4, 239. 247. 202. 397.
403
australis, Harpinia. 535
Harpiniopiis, 5)5
Pomharpinia. 190
Protophoxui, 145, 176, 190
Aviccnnia marina, 30
axutlaiita, Harpinia. 534
Pseudharpinia, 5)4
babaneekus, Birubius, 19, 176. 196. 198. 199, 200. 201. 276,
281. 285. 288. 292. 343
barnardi, Pontharpinia. 145
bassi, Phoxocephalus, 418, 422, 427. 441, 442, 443
Phoxus, 418
Batuto, 21. 91. 147. 155, 167, 175, 529, 5)0, 531
stimpsoni, 145. 146. 531
bateiT, Birubius, 145, 161, 197. 199, 200, 201, 222, 298, 309,
312, 317, 320. 347, 351
Phoxus 145, 176, 190, 312
batturi, Kotla, 161
bicuspidatus, Paraphoxus, 39, 145
bidentata, Harpinia, 536
birjulini, Harpinia, 534
Pseudharpinia, 5)4
birubi, Tickalerus, 21. 397, 398, 408. 414
Birubius, 7, 9. 12, 17, 18, 19, 20, 21, 22, 23, 24, 25, 27, 29, 32,
39, 88, 147, 155, 175, 176, 190, 191, 192, 193, 199, 207. 251,
275, 298, 337, 338, 351, 356, 375, 389, 397, 403, 415
apart, 196, 198, 200, 201, 228, 234, 239, 247, 292, 397, 403
babaneekus, 19, 176, 1%, 198, 199, 200, 201, 276, 281, 285,
288, 292. 343
bateiT, 145, 161, 197, 199, 200, 201, 222, 298, 309, 312, 317,
320, 347, 351
booleus, 21. 92. 93. 1%, 198, 200, 201, 272, 275, 365
cartoo, 1%, 198, 199, 201, 234, 239, 242, 247, 251
chintoo, 196, 198, 200, 201. 264, 268, 271. 272, 275, 343, 365
eake, 197. 199, 200, 201, 292, 356. 357, 362, 365. 370
eleebanus. 19. 176. 197. 198. 200, 201, 212, 281. 337, 338,
343. 362, 370. 388
gallangus, 196. 198. 200, 201, 212, 239, 247, 251, 343, 362,
365. 370, 389
gambodeni, 196. 200, 201, 207, 222, 292, 297, 298, 304, 309,
320, 325. 329, 362, 397
gelarus, 196, 198, 199. 200. 201, 281, 288
jirrandus, 197. 199, 200, 201. 222, 298, 312, 318, 320. 343,
347, 351, 355. 356
kabbulinus, 197, 199. 200, 201, 356, 357, 362, 365, 370
kareus, 1%, 198, 200, 222, 227, 247, 251
karobrani, 176, 196, 198, 200, 201, 268, 271, 272, 281, 338,
343, 370
kinkus, 196, 197, 199, 200, 201, 298, 325, 329, 334
kokorus, 197. 199. 200. 201, 298, 320, 325, 329, 334
kyeemus, 197, 199, 200, 201. 298, 309, 312, 329, 334
lorus, 24, 194. 198, 200, 201, 207, 212, 215, 247, 251, 343, 389
lowannus, 197, 199, 200, 222, 298, 304, 309, 312, 320, 334
maamus, 18, 197, 199. 200, 201, 298. 299, 325, 329
maldus, 197, 199, 200, 201, 370, 375, 380
mayamayi, 9, 87, 88, 92, 93. 152, 196, 198, 200, 201. 207, 251,
260, 264, 266, 272, 275, 338, 343, 370
muldarpus, 196, 198, 200, 201, 222, 239, 242, 247, 251, 365,
370, 389
munggai, 197, 199. 200, 201, 222, 298, 309, 329, 334, 362, 370
myallus, 196, 198, 200, 201, 216, 222, 227, 234, 239, 247, 251,
298, 351, 355, 362
nammuldus, 21, 196, 198, 199, 200, 201, 212, 222, 247, 343,
389
narus, 197, 199, 200, 201, 288, 292, 343, 362, 370
panamunus, 192. 193. 194, 198, 199, 200, 201, 207, 212, 222,
234, 239, 247, 251, 260, 281, 292
quearus, 196, 198, 199, 200, 201, 281, 285
rostratus, 146, 351
taldeus, 21, 24, 197, 199, 200, 201, 292, 355, 356, 362, 365,
370, 375, 380, 388
thalmus, 196, 198, 199, 201, 2)9, 242, 247
ularitus, 93, 197, 198, 200, 201, 334, 338, 343, 370
wirakus, 196, 198, 200, 201, 260, 264, 268, 272, 275
wulgaru, 17, 19, 197, 199, 200, 201, 338, 380, 388
yandus, 197, 199, 200, 201, 370, 375, 380
yorlunus, 197, 198, 199, 200, 201, 247, 351, 355, 356, 357,
362, 365, 370
545
546 SMITHSONIAN CONTRIBUTIONS TO ZOOI OCY
species, 312
booleus, Birubius, 21, 92, 93, 196, 198, 200. 201, 272, 275, 565
Booranus, 13, 18, 21, 46, 47, 48, 49, 70, 71, 72, 87
tikeri, 71,72, 76, 85
wangoorus, 71, 72, &/, 85
weemus, 71, 72, 80, 85
borra/tw, Kulgaphoxus, 403, -/W, 408, 414
brevidactylus, Metaphoxus, 469
Vasco, 469
brevirostris, Harpinia, 534
Pseudharpinia, 534
Brolgus, 25, 87, 88, 89, 90, 92, 96, 104, 116, 119, 125, 130, 134,
145, 167, 192, 365, 488
koongarrus, 97, 98, 108,116, 118, 119
mahmak, 97, 98, 103,108, 112, 116, 118, 119, 130
millinus, 97, 98, 103,104, 108, 112, 116, 118,119, 130, 133
tattersalli, 9, 87, 97, 98, 103, 108, 112, 116, 143, 146, 176
tavelus, 97, 98,108,113, 116, 117, 118, 143
burleus, Phoxocephalus, 418, 435, 439, 441
bychovskii, Joubinella, 153
cabotensis, Harpinia, 536
cadgeeus, Kulgaphoxus, 167, 397, 403, 408, 414
calcarata, Pontharpinia, 145
capensis, Harpinia laevis, 535
Harpiniopsis, 535
capillatus, Trichophoxus, 24, 86, 145
cariniceps, Harpinia 534
Pseudharpinia, 534
cartoo, Birubius, 196, 198, 199, 201, 234, 239, 242, 247, 251
centralis, Pontharpinia, 145
chelatus, Paraphoxus, 33,145
Phoxocephalus, 465
chintoo, Birubius, 196, 198, 200, 201, 264, 268, 271, 272, 275,
343, 365
ciliata, Joubinella, 153, 154
cmca, Harpinia, 534
Pseudharpinia, 534
cognatus, Paraphoxus, 32, 145
Compactogammarus, 34
cornuta, Metharpinia, 24, 32, 145, 416
coxalis, Coxophoxus, 530
Phoxocephalus, 530
Coxophoxus, 6, 32, 529, 530
coxalis, 530
hidalgo, 530
crenulata, Harpinia, 536
crenuloides, Harpinia, 536
Cunmurra, 87, 88, 89, 92, 93, 152, 191, 192, 275, 365
itickerus, 92,93, 275, 338, 365
curtipes, Harpinia, 536
daboius, Paraphoxus, 32, 145
dellavallei, Harpinia, 536
dentata, Pseudharpinia, 534
dinjerrus, Tipimegus, 47, 49, 55, 58, 70, 71
Diogodias, 416, 417, 467, 468
littoralis, 468
longicarpus, 468
platyrostris, 468
take, Birubius. 197. 199. 200, 201. 292. 3%. ?57, 362. 365. 570
eleebanus, Birubius. 19, 176, 197. 198, 200, 201. 212. 281. 337,
338, 343, 362. 370. 388
Elpeddo, 87. 88, 89. 90. 118, 119. 134. 144, 145
kaikai, 18, 119, 144
emeryi, Harpiniopsis, 5)5
Entlevoura, 22
episloma, Pontharpinia. 145
epistomata, Harpiniopsis, 535
epistomus, Paraphoxus, S3
errichus, Yan, 389, 390. 393
excavata, Harpinia, 534
Pseudharpinia, 534
falcatus, Leptophoxus, 448
Phoxus, 448
falcatus falcatus, Leptophoxus, 448
falcatus icelus, Leptophoxus, 448
Falklandella. 21
fatigans, Paraphoxus, 32. 145
floridana. Pontharpinia. 145
florkl.'inus. Paraphoxus, 32
frequens, Metaphoxus, 449, 456, 460, 465
fuegiensis, Paraphoxus. 144
Parharpinia. 134. 144. 145
WildusT, 18, 32, 134, 144, 145
fulgens, Harpiniopsis, 535
fultoni, Metaphoxus, 449, 465, 466. 475
Phoxocephalus, 449, 465
galera, Harpiniopsis, 535
gallangus, Birubius, 196, 198. 200, 201. 212, 239. 247, 251, 343,
362, 365, 370, 389
gambodeni, Birubius, 196, 200, 201, 207. 222, 292, 297. 298.
304, 309, 320, 325, 329, 362, 397
Gammarus, 22
Ganba, 87, 88, 90, 97, 119,124,125, 130, 134, 145
pellati, 125
gelarus, Birubius, 196, 198, 199, 200, 201, 281, 288
gemmatus, Paraphoxus, 145
geniculatus, Phoxus, 145
gilesi, Mandibulophoxus, 90, 91
grandis, Phoxus, 145
gurjanovae, Harpinia, 535
Harpiniopsis, 535
Harpinia, 6, 18, 33, 37, 528, 529, 535, 536
abyssalis, 534
abyssi, 536
amundseni, 535
antennaria, 536
australis, 535
ayutlanta, 534
bidentata, 536
birjulini, 534
brevirostris, 534
cabotensis, 536
NUMBER 245 547
cariniccps, 534
dnca, 534
crenulata, 536
crenuloidet, 5)6
curtipes, 5)6
dellavallei, 5)6
excavate. 534
gurjanovae. 535
kobjakovae, 535
laevis, 535, 5)6
laevis capensis, 535
latipes. 534
miharamsis, 535
moisecvi, 535
mucronala, 5)6
nadania, 535
neglecta, 536
obi uiifrom, 534
oculata. 533
ophtbalmica, 533
orirmalit. 535
paciftca. 535
palabria. 5)4
pectinala, 5)6
plumoia, 5)6
propinqua, 5)6
salebroia, 535
ichurini, 535
serrate, 5)6
spaercki. 534. 535
taratovi, 535
truncate, 5)6
vallini. 534
wandichia, 535
Harpiniopsis, 6, 529, 530. 532. 5)4, 536
amundseni, 5)5
aus traits, 535
caperuis, 535
emeryi, 535
epistomata, 535
fulgens, 535
galera, 535
gurjanovae, 535
kobjakovae, 5)5
miharaensis, 535
moiseevi, 535
nadania, 535
naiadis, 535
orientalis, 535
pacifica, 535
percellaris, 535
petulans, 535
profundis, 535
salebrosa, 535
sanpedroensis, 534
schurini, 535
similis, 535
spaercki, 535
tarasovi, 535
triplex, 535
wandichia, 535
species D, 535
heterocuspidatus, Paraphoxus, 145
Heterophoxus, 1, 6. 17, 23, 32, 272. 529, 532
oculatus, 17, 32, 533
ophthalmica, 53)
pennatus, 533
stephenseni, 532
trichosus, 533
videns, 533
Heterozostera tasmanica, 30
hidalgo, Coxophoxus, 530
Holbolli, Phoxocephalus, 17, 33. 441
Phoxus. 418. 441
homilis, Phoxocephalus, 418, 431, 441
Hopiphoxus, 416, 417. 469
simillimus, 470
hurleyi, Proharpinia, 532
homilis, Phoxocephalus, 418, 451,441
icelus, Leptophoxus falcatus, 448
intermedia, Pontharpinia, 145, 531
itickerus, Cunmurra, 92. 93, 275, 338, 365
Japara, 17, 18. 20. 23. 25, 416. 474, 475, 481
papporus, 475
Jerilderia, 35, 416. 442, 443
joubiphoxus, 442, 443
jhrandus, Birubius, 197, 199. 200, 201, 222, 298, 312, 318. 320,
343, 347. 351.355, 356
jonesi, Paraphoxus, 145
Joubinella, 18. 22, 29, 33, 35, 39, 152,153, 441. 442, 443, 481
bychovskii, 153
ciliata, 153, 154
strelkovi, 153, 154
traditor, 153, 154
tzuetkovae, 153
species, 154
joubiphoxous, Jerildaria, 442, 443
kabbulinus, Birubius, 197, 199, 200, 201, 356, 357 362, 365,
370
kaikai, Elpeddo, 18, 119, 144
kalduhe, Limnoporeia, 488, 517, 521
kalkro, Tipimegus, 47, 49, 64, 70
kangulun, Tipimegus, 47, 49, 52, 58, 64, 70
kareus, Birubius, 196, 198, 200. 222, 227, 247, 251
karobrani, Birubius, 176, 196, 196; 200, 201, 268, 271, 272, 281,
338, 343, 370
keppeli, Phoxocephalus, 418, 422, 441
kergueleni, Phoxocephalus, 427, 431, 441, 442
Phoxus, 418, 441
kingi, Limnoporeia, 31, 488, 489, 501
kinkus, Birubius, 196, 197, 199, 200, 201, 298, 325, 329, 334
kobjakovae, Harpinia, 535
Harpiniopsis, 535
kokorus, Birubius, 197, 199, 200, 201, 298, 320, 325, 329, 334
548
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Kondoleus, 17, 18, 21, 24, 25, 416, 475, 480, 481, 506
tekin, 481
koongarrus, Brolgus, 97, 98, 108, 116, 118, 119
Kotla, 17, 19, 21, 25, 147, 152, 161, 167
batturi, 161
Kroyerii, Phoxus, 465
kukathus, Phoxocephalus, 418. 422, 427, 4S1
Kulgaphoxus, 24, 25, 191, 397, 402, 403, 404
borralus, 403, 404, 408, 414
cadgeeus, 167, 397, 403, 408, 414
kunarella, Yammacoona, 167
Kuritus, 20, 87, 88, 90, 119, 125, 729, ISO, 134, 145
nacoomus, 130
kyeemus, Birubius, 197, 199, 200, 201, 298, 309, 312. 329, 334
laevis, Harpinia, 535, 536
laevis capensis, Harpinia, 535
latipes, Harpinia, 534
Pseudharpinia, 534
lea, Rikkarus, 470, 471
Leongathus, 19, 146, 147, 176
nootoo, 147
Leptophoxoides, 35, 41, 416, 446
molaris, 447
Leptophoxus, 20, 416, 447, 475
falcatus falcatus, 448
falcatus icelus, 448
uncirostratus, 91
Limnoporeia, 6, 21, 34, 416, 417, 4S\,487, 488, 522
kalduke, 488, 517, 521
kingi, 31,488, *S9, 501
maranowe, 488, 489, 493, 501, 506, 511
ungamale, 488, 506, 511, 517, 521
wakkine, 488, 577, 517, 521
woorake, 488, 507, 511
yarrague, 488, 489, 495, 501, 506, 511, 521
littoralis, Diogodias, 468
Metaphoxus, 468
longicarpus, Diogodias, 468
Metaphoxus. 468
longirostris, Metharpinia, 32, 145, 416
Paraphoxus, 416
lorus, Birubius, 24, 194, 198, 200, 201, 207, 212, 215, 247, 251,
343, 389
lowannus, Birubius, 197, 199, 200, 222, 298, 304, 309, 312, 320,
334
lucubrans, Paraphoxus, 145
maamus, Birubius, 18, 197, 199, 200, 201, 298, 299, 325, 329
maculatus, Paraphoxus, 145
Phoxus, 145
mahmak, Brolgus, 97, 98, 103,108, 112, 116, 118, 119, 130
maldus, Birubius, 197,199, 200, 201, 370, 375, 380
Mandibulophoxus, 18, 23, 32, 87, 89, 90, 91, 147, 155, 167, 175,
531
gilesi, 90, 91
stimpsoni, 531
uncirostratus, 91, 146
maranowe, Limnoporeia, 488, 489, 493, 501, 506, 511
marina, Avicennia, SO
Matong, 25. 146, 147. 152. 75*, 155, 161. 167
matong, 146, 755, 312
matong, Matong, 146, 755, 312
maxima, Pontharpinia, 145
mayamayi. Birubius, 9. 87. 88. 92. 93. 152. 1%. 198. 200. 201.
207. 251. 260, 264. 266. 272. 275. 338. 343. 370
Metaphoxoides, 6, 416, 417. 466, 467
picardi, 466, 467
zavorus, 466. 467
Metaphoxus, 6. 33, 89. 416, 417. 448, 449. 468. 469. 470. 475.
481.488
brevidactylus. 469
frequens, 449. 456, 460. 465
fultoni, 449. 465, 466, 475
littoralis. 468
longicarpus, 468
mintus, 449. 460, 465
pectinatus, 449, 463
platyrostris. 468
simillimus. 470
tuckatuck, 449, 456. 460. 465
tulearensis, 449, 466
typicus. 449. 465
yaranellus, 449. 456, 460
zavorus, 466
species. 466, 467
Metharpinia, 144. 191, 415, 416
cornuta. 24, 32, 145, 416
longirostris, 32, 145, 416
Microphoxus, 21, 32. 191, 414, 416
minimus, 29, 415
miharaensis, Harpinia, 535
Harpiniopsis, 535
milleri, Pontharpinia, 7*5
millinus, Brolgus, 97, 98, 103.104, 108, 112, 116, 118, 119, ISO,
13S
minimus, Microphoxus, 29, 415
mintus, Metaphoxus, 449, 460, 465
moiseevi, Harpinia, 535
Harpiniopsis, 535
molaris, Leptophoxoides, 447
mucronata, Harpinia, 536
muelleri, Zostera, 30
muldarpus, Birubius, 196, 198, 200, 201, 222, 239, 242, 247,
251, 365, 370, 389
mullokus, Wildus, 134, 138, 143
munggai, Birubius, 197, 199, 200, 201, 222, 298, 309, 329, 334,
362, 370
myallus, Birubius, 196, 198, 200, 201, 216, 222, 227, 234, 239,
247, 251, 298, 351, 355, 362
nacoomus, Kuritus, 130
nadania, Harpinia, 535
Harpiniopsis, 535
naiadis, Harpiniopsis, 535
nammuldus, Birubius, 21, 196, 198, 199, 200, 201, 272, 222,
247, 343, 389
narus, Birubius, 197, 199, 200, 201, 288, 292, 343, 362, 370
NUMBER 245 549
nasuta, Pontarpinia [sic], 145
nrglecta. Harpinia. 536
Niphargogammarus. 34, 35
N'iphargoides, 34
nootoo, Leongathus, 147
Obesogammarus, 34
obliqua, Parharpinia. 32, 145
nlnusidrns, Pontharpinia, 145
obtusifrons, Harpinia, 534
Pseudharphiia, 534
oblmus. Phoxui, 146
ochotica. Pararpinia [sic], 146
oculata. Harpinia, 533
oculalus. Heterophoxus, 17, 32, 533
Paraphoxm, 16. 31, 145
Phoxut. 145
ophlhalmica, Harpinia, 533
Hrterophoxm, 533
oriental!*. Harpinia. 535
Harpiniopsis, 535
pacifica, Harpinia, 535
Harpiniop$i$, 535
1>annmumt%. Birubim. 192. 193. 194, 198, 199, 200. 201, 207.
212. 222. 234. 239. 247. 251. 260. 281. 292
palabria, Harpinia. 534
papporus, Japara, 475
Paradotaminc, 7
Parandaniexis, 22
Paraniphargoidcs, 34
Paraphoxm, 6. 7. 17. 21, 87, 89, ISO. 144, 145, 365
abroniui, 145
bicmpidatus. 39. 145
chelatus. 33. 145
cognatus. 32, 145
daboius. 32, 145
epistomus, 33
fatigans, 32. 145
floridanus. 32
fuegiensis, 144
gemmatus, 145
heterocuspidatus, 145
jonesi. 145
longirostris, 416
lucu brans, 145
macula tus, 145
oculatus, 16, 31, 145
pinguis, 41
pyripes, 33, 145, 146
rakiura, 87, 146
robustus, 146
rostrata(us). 146
similis, 146
? simplex, 145
spinibasus, 33, 146
spinosus, 16, S3, 146
stebbingi, 47, 70, 146
stenodes, 146
subuncigerus, 146
tattcrsalli, 97, 98, 146
uncirostrata, 91
variatus, 35, 146
vigitegus, 24. 146
villosus, 176
waipiro, 134, 144, 146
sp.. 146, 155
Pararpinia [sic] ochotica, 146
simplex, 145
Parharpinia, 17, 33, 144, 175, 176. 190, 192, 415
fuegiensis, 134, 144, 145
obliqua, 32, 145
rotundifrons, 32, 146
simplex, 145
sinuata, 32, 146
sinuatut, 176
uncigera, 146
villosa, 9, 25, 98, 146, 176, 189, 272, S4S
wane, 175, 176, 185, 189
species, 176
pectinata, Harpinia, 536
pectinatus, Metaphoxus, 449, 465
Phoxocephalus, 449, 465
pellati, Ganba, 125
pen na tus, Heterophoxus, 533
percellaris, Harpiniopsis, 535
petulans, Harpiniopsis, 535
Phoxocephalus, 19, 33, 35. 39, 41. 89, 416, 417, 418, 4S1, 441
bassi, 418, 422. 427, 441, 442. 44S
burleus, 418, 435, 439, 441
chelatus, 465
coxalis, 530
fukoni, 449. 465
holbolli, 17, SS, 441
homilis, 418, 431, 441
keppeli, 418, 422, 441
kergueleni, 427, 431, HI, 442
kukathus, 418, 422, 427, 431
pectinatus, 449, 465
regium, 418, 427, 435, 439, 441. 442
rupullus, 418. 431, 435, 439, 441
simplex, 465
tenuipes, 418, 442
tunggeus, 418, 427, 431
species, 441
Phoxus, 417
bassi, 418
batei, 145, 176, 190, 312
ralcatus, 448
geniculatus, 145
grandis, 145
Holbolli, 418, 441
kergueleni. 418. 441
Kroyerii, 465
macula tus, 145
obtusus, 146
oculatus, 145
plumosus, 5S6
550 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
simplex, 465
unicirostratus, 90, 91, 146
villosus, 146, 175, 176
picardi, Metaphoxoides, 466, 467
pillare, Uldanamia, 488, 521, 522
pinguis, Paraphoxus, 41
Pontharpinia, 41, 46, 70, 146
Urothoe, 41, 146
platyrostris, Diogodias, 468
Metaphoxus, 468
plumosa, Harpina, 536
plumosus, Phoxus, 536
Pontarpinia [sic] nasuta, 145
Pontharpinia, 17, 18, 19, 20, 21, 23, 24, 25, 35, 37, 40, 41, 47,
48, 70, 90, 91, 144, 147, 155, 167, 192. 193, 207, 443, 531
austral is, 190
barnardi, 145
calcarata, 145
central is, 145
epistoma, 145
floridana, 145
intermedia, 145, 531
maxima, 145
milleri, 145
obtusidens, 145
pinguis, 41, 46, 70, 146
rostrata(us), 146, 312
stimpsoni, 91, 146. 531
tridentata, 146
uncinata, 32, 146
uncirostratus, 91
villosa, 176
sp., 146, 176
Pontogammarus, 34
profundis, Harpiniopsis, 535
Proharpinia, 1, 32, 33, 529, 531
antipoda, 532
hurleyi, 532
stephenseni, 532
tropicana, 532
propinqua, Harpinia, 536
Protophoxus, 25, 33, 144, 175, 189, 190, 192, 415
australis, 145, 176, 190
Pseudharpinia, 1, 17, 529, 530, 533, 535
abyssalis, 534
ayutlanta, 534
birjulmi, 534
breuirostris, 534
cariniceps, 534
cinca, 534
dentata, 534
excavata, 534
latipes, 534
obtusijrons, 534
vallini, 534
pyripes, Paraphoxus, 33, 145, 146
quearus, Birubius, 196, 198, 199, 200, 201, 281, 285
rakiura, Paraphoxus, 87, 146
Waitangi, 33, 87. 146
regium, Phoxoccphalus, 418, 427, 435, 439, 441, 442
Rikkarus, 416. 417. 470
lea. 470, 471
robustus. Paraphoxus, 146
rostrata(us), Pontharpinia. 146, 312
rostratus(a), Birubius, 146, 351
Paraphoxus, 146
Urothoe. 146
rotundifrons, Parharpinia, 32. 146
rupullus, Phoxoccphalus, 418, 431. 435. 439. 441
salebrosa, Harpinia, 535
Harpiniopsis, 535
sanpedroensis, Harpiniopsis. 534
schurini, Harpinia, 535
Harpiniopsis, 535
serrata, Harpinia, 536
similis, Harpiniopsis, 535
Paraphoxus, 146
simillimus, Hopiphoxus, 470
Metaphoxus, 470
simplex, Paraphoxus?, 145
Pararpinia [sic], 145
Parharpinia, 145
Phoxocephalus, 465
Phoxus, 465
sinuata, Parharpinia, 32, 146
sinualus, Parharpinia. 116
sanpedroensis, Harpiniopsis, 534
spaercki, Harpinia, 534, 535
Harpiniopsis, 535
sp., Paraphoxus, 146, 155
Pontharpinia, 146, 176
species, Joubinella, 154
Metaphoxus, 466, 467
Parharpinia, 176
Phoxocephalus, 441
D, Harpiniopsis, 535
4N, Tipimegus, 70
5N, Tipimegus, 70
spinibasus, Paraphoxus. 33, 146
spinosus, Paraphoxus, 16, 33, 146
stebbingi, Paraphoxus, 47, 70, 146
Tipimegus, 41, 70, 146
stenodes, Paraphoxus, 146
stephenseni, Heterophoxus, 532
Proharpinia, 532
stimpsoni, Basuto, 145, 146, 531
Mandibulophoxus, 531
Pontharpinia, 91, 146, 531
strelkovi, Joubinella, 153, 154
subuncigerus, Paraphoxus, 146
Syndexamine, 506
taldeus, Birubius, 21, 24, 197, 199, 200, 201, 292, 355, 356, 362,
365, 370, 375, 380, 388
NUMBER 245 551
tarasovi, Harpinia, 5S5
Harpiniopsis, 5)5
tasnunica, Hetcrozostera, 30
tattenalli, Brolgus, 9, 87, 97. 98, 103, 108. 112, 116, 143, 146.
176
Paraphoxus. 97. 98, 146
tavelus, Brotgus. 97. 98. 108.11), 116, 117, 118, 143
tekin, Kondoleus, 481
tetiuipes, Phoxocephalus, 418, 442
thalerus, Tipimegus, 47. 49, 55, 57. 58, 64, 70, 71
thalmus, Birubius, 196. 198. 199, 201, 237, 2)9, 242. 247
thambaroo, Wildus, 1)4, 143
Tickalerus. 24. 191. )96. 403. 404
birubi, 21. 397. )98, 408. 414
tiendi, Yan, 389. 390, 396
tikeri, Booranus, 71. 72. 76, 85
Tipimegus, 18. 20, 21. 23. 24. 25, S3. 46. 47, 48. 49. 55, 72. 86,
87. 192, 415
dinjerrus. 47. 49. 55, 58, 70. 71
kalkro, 47, 49, 64, 70
kangulun. 47. 49. 52. 58, 64, 70
ttebbingi. 41. 70, 146
f/wterw. 47. *9, 35. 57. 58. 64. 70. 71
specie* 4N, 70
?peciet 5N. 70
traditor, Joubinella, 15), 154
Trichophoxus, 17. 33. 46. 47. 48, 49, 85, 86. 87. 144, 192
capUlatus, 24. 86, 145
trichosus, Heterophoxus, 53)
tridentata, Poniharpinia, 146
triplex, Harpiniopsis, 535
Trischizostotna, 22
tropicana, Proharpinia, 5)2
truncata, Harpinia, 536
tuckatuck, Metaphoxus, 449, 456. 460, 465
tulearensis, Metaphoxus, 449, 466
tunggeus, Phoxocephalus, 418, 427, 431
typicut, Metaphoxus, 449, 465
tzvetkovae, Joubinella, 153
ularitus, Birubius, 93.197,196, 200, 201,3)4, 338. 343. 370
Uldanamia, 21, 416, 417, 488. 522
pUlare, 488, 521, 522
uncigera, Parharpinia, 146
uncinata, Pontharpinia, 32, 146
undrottrata, Paraphoxus, 91
uncirostratus, Mandibulophoxits, 91, 146
Phoxus. 90, 91, 146
Pontharpinia, 91
ungamale, Limnoporeia, 488, 506, 511.517,521
Uroniphaigoides, 34
Urothoe, 36
pinguis, 41, 146
rostra tus, 146
vallini, Harpinia, 534
Pseudharpinia, 5)4
variatus, Paraphoxus, 35, 146
Vasco. 416. 417. 468
brevidactylus, 469
videns, Heterophoxus, 5))
vigitegus, Paraphoxus, 24,146
villosa, Parharpinia, 9, 25, 98. 146,176, 189. 272. 343
Pontharpinia, 176
villosus, Paraphoxus, 176
Phoxus, 146, 175, 176
waipiro, Paraphoxus, 134, 144, 146
Wildus?, S3. 134.144, 146
Waitangi, 24, 46, 47. 48. 86, 87
rakiura, 33, 87. 146
wakkine, Limnoporeia, 488. 501, 511
wandichia, Harpinia, 535
Harpiniopsis, 5)5
wangoorus, Booranus, 71, 72, 81, 85
warte, Parharpinia, 175, 176. 185, 189
weemus, Booranus, 71, 72, 80, 85
Wildus. 87, 88, 89. 90, 119. 125. 130,133.134, 144, 145
f fuegiensis, 18, 32, 134,144,145
mullokus, 134,1)8, 143
thambaroo, 134, 143
T waipiro. 33, 134,144, 146
wirakus, Birubius. 196, 198, 200, 201,260, 264, 268, 272. 275
woorake, Limnoporeia, 488,501, 511
wulgaru, Birubius, 17,19,197,199, 200, 201, 338.380, 388
Yammacoona, 17,18, 20. 147, 152,166, 167
kunarella, 161
Yan, 23,191, 388, 389. 403, 404
errichus, 389, 390. 393
tiendi, 389, 390, 396
yandus, Birubius. 197,199, 200, 201,370. 375. 380
yaranellus, Metaphoxus, 449. 456, 460
yarrague, Limnoporeia, 488, 489, 495. 501, 506. 511, 521
yorlunus, Birubius. 197,198, 199, 200, 201. 247.351, 355, 356,
357, 362, 365, 370
zavorus, Metaphoxoides, 466, 467
Metaphoxus, 466
Zobracho, 35
Zostera mudleri, 30

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