Gammaridean Amphipoda
of Australia, Part V:
Superfamily Haustorioidea
J. LAURENS BARNARD
and
MARGARET M. DRUMMOND
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 360
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 3 6 0
Gammaridean Amphipoda
of Australia, Part V:
Superfamily Haustorioidea
J. Laurens Barnard
and Margaret M. Drummond
SMITHSONIAN INSTITUTION PRESS
City of Washington
1982
A B S T R A C T
Barnard, J. Laurens, and Margaret M. Drummond. Gammaridean Amphi-
poda of Australia, Part V: Superfamily Haustorioidea. Smithsonian Contributions
to Zoology, number 360, 148 pages, 58 figures, 1 table, 1982.?New genera and
species in the superfamily Haustorioidea are described. The Haustorioidea are
reorganized by rejection of the Pontoporeiidae and the description of new
families Urohaustoriidae, Zobrachoidae, and Condukiidae; the Haustoriidae,
Phoxocephalidae, and Platyischnopidae are rediagnosed to fit this haustorioid
pattern. The described species of Urohaustorius are clarified (9 new species) and
the new genera Gheegerus (1 new species), Narunius (1), Tuldarus (2), Tottungus
(1) and Dirimus (1), with affinities to Urohaustorius, are established. The
Zobrachoidae are established to contain Zobracho and the new genera Bumeralius
(1) and Prantinus (1). The new genus Condukms (1) is assigned to a new family
of its own. The Urothoidae are defined to include Urothoe, Urothoides, Carangolia,
and Cumcus.
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 DESIGN: The coral Montastrea
cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data
Barnard, Jerry' Laurens, 1928-
Gammaridean Amphipoda of Australia.
(Smithsonian contributions to zoology ; no. 103. 139, 245)
Vol. 3 also has special title: The Phoxocephalidae.
Vol. 5 also has special title: Superfamily Haustorioidea.
Vol. 3, by J. L. Barnard and M. M. Drummond.
Bibliography: p.
Includes bibliographies.
Supt. of Docs, no.: SI 1.27:103
1. Amphipoda?Australia. I. Drummond, Margaret M., joint author. II. Title. III. Series:
Smithsonian Institution. Smithsonian contributions to zoology, no. 103 [etc.]
QL1.S54 no. 103. etc. |QL444.A5] 591s [595.3'71] 72-601600 AACR1
Contents
Page
Introduction 1
Acknowledgments 1
Methods 2
Organization of Illustrations 3
Definition of Terms 3
Text Abbreviations 4
Figure Abbreviations 4
Superfamily HAUSTORIOIDEA Stebbing, 1906 4
Classification in Haustorioidea 5
Evolutionary Pattern in Haustorioidea 7
Key to Families of Haustorioidea 12
PHOXOCEPHALIDAE 12
PLATYISCHNOPIDAE 13
UROTHOIDAE 14
Key to Genera of Urothoidae 15
PHOXOCEPHALOPSIDAE 15
Phoxocephalopsis Schellenberg 16
Phoxocephalopsis zimmen Schellenberg 16
ZOBRACHOIDAE, new family 18
Key to Genera of Zobrachoidae 18
Zobracho J.L. Barnard 19
Zobracho canguro J.L. Barnard 20
Bumeralius, new genus 27
Bumeralius buchahus, new species 28
Prantinus, new genus 36
Prantinus talanggi, new species 38
UROHAUSTORIIDAE, new family 45
Key to Genera of Urohaustoriidae 47
Urohaustonus Sheard 47
Key I to Selected Species of Adult Urohaustorius 50
Key II to Selected Species of Adult Urohaustorius 51
Key III to Selected Species of Adult Urohaustorius 52
Urohaustorius pulcus, new species 53
Urohaustorius pulcus, morph 872 59
Urohaustorius pentinus, new species 61
Urohaustorius merkanius, new species 64
Urohaustonus wingaro, new species 70
Urohaustorius parnggius, new species 74
Urohaustorius yurrus, new species 76
Urohaustonus urungan, new species 80
Urohaustorius species M 84
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
I rohaustonus perkeus, new species 85
Urohaustonus halei Sheard 89
Urohaustonus vercoi Sheard 97
Urohaustonus metungi Fearn-Wannan 97
Urohaustonus gunm, new species 103
Gheegerus, new genus 106
Gheegerus garbaius, new species 108
Narunius, new genus I l l
Narumus tallerkus, new species 112
Tuldarus, new genus 118
Key to Species of Tuldarus 119
Tuldarus cangellus, new species 119
Tuldarus bannius, new species 123
Tottungus, new genus 126
Tottungus tungus, new species 127
Dmmus, new genus 132
Dinmus tarlitus, new species 133
HAUSTORIIDAE 136
CONDUKIIDAE, new family 138
Condukius, new genus 139
Condukius karkan, new species 139
Literature Cited 146
Index 147
Gammaridean Amphipoda of Australia,
Part V: Superfamily Haustorioidea
J. Laurens Barnard
and Margaret M. Drummond
Introduction
Marine taxa of the families Urohaustoriidae,
Zobrachoidae, and Condukiidae are treated
herein. This continues our studies of those gam-
maridean groups in Australia needing report most
urgently because of their value to other biologists
involved in impact surveys, monitoring programs,
and population ecology.
The haustorioid groups are especially impor-
tant because of their diversity, abundance of
individuals and obscure taxonomy. As in parts
III and IV the bulk of our material originates
from the various surveys conducted by the Ma-
rine Studies Group in Melbourne, but an increas-
ing percentage of specimens is coming from other
institutions launching major studies of the marine
environment in New South Wales and Queens-
land.
ACKNOWLEDGMENTS.?The first author is espe-
cially indebted to Dr. F.H. Drummond, The
University of Melbourne, for his gracious hospi-
tality and many kindnesses to him during stays
in Melbourne in 1971, 1976, and 1980, helping
with accomodations, field trips and valuable in-
formation; gratitude must be expressed also to:
Dr. and Mrs. Jerry D. Kudenov of the Marine
/ . Laurens Barnard, Department of Invertebrate Zoology, National
Museum of Natural History, Smithsonian Institution, Washington,
D. C. 20560. Margaret M. Drummond, National Museum of I ictona,
71 Victoria Crescent, Abbotsford, Victoria 3067 Australia.
Pollutions Study group, and their colleagues, Dr.
Gary B. Poore, Dr. and Mrs. Alistair J. Gilmour
for their hospitality and valuable aid; also to Dr.
Murray Littlejohn, Mr. John Dorsey and Ms.
Michelle Barson of the University of Melbourne:
to Dr. Harold Heatwole of the University of New-
England, Armidale; to Dr. Leon Collett and Mr.
Phillip J. Gibbs of New South Wales Fisheries
Dept.; and Mr. Terry Walker of the University
of Tasmania, all for their help with logistics or
materials for our studies.
At the Western Australian Museum, the first
author must again thank Mrs. Shirley M. Slack-
Smith, Dr. Ray George, Dr. Barry R. Wilson,
Director, National Museum of Victoria, Loisette
Matilda Marsh and Dr. Fred E. Wells for their
hospitality and assistance with logistics and mu-
seum materials during three trips there in 1968
and 1976.
Travel to Australia for the first author was
supported by a research award from the Secretary
of the Smithsonian Institution, S. Dillon Ripley;
funds for inking the illustrations came from the
Smithsonian Research Foundation to support
Carolyn L. Cox and Irene Y. Jewett in this task.
Dr. W. Duane Hope, Chairman of Invertebrate
Zoology, was of great assistance in solving diffi-
cult logistical problems. Mr. Roland H. Brown
and Mrs. Dessie M. Best, Janice Clark and Eliz-
abeth B. Harrison also assisted the first author.
1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
At the Australian Museum in Sydney a debt of
gratitude is owed to Director, Dr. DJ.G. Griffin,
and curators, Dr. James K. Lowry and Dr. Patri-
cia Hutchings for their valuable assistance during
visits there in 1968, 1971, 1976, and 1980.
In Brisbane, Prof. W. Stephenson of the Uni-
versity of Queensland gave full access to his val-
uable material, laboratory and hospitality, and
Mr. Ronald Munroe of the Queensland Museum
provided access to other valuable specimens.
Prof. W.D. Williams of the University of Ade-
laide gave graciously of his expert advice and fine
hospitality in Adelaide, as he has on so many
occasions in the past. Finally, Charline M. Bar-
nard continued her valuable assistance in man-
aging many aspects of the laboratory work, logis-
tics, and research documentation.
This paper is listed as publication No. 314 in
The Environmental Series of the Ministry for
Conservation in Victoria.
Most of the materials used in this study have
come from benthic surveys in Western Port
(1964-1965 and 1973-1974) and in Port Phillip
Bay (1971-1973). Acknowledgment and appre-
ciation to organizations and individuals have
been expressed more fully in a previous publica-
tion (Barnard and Drummond, 1978).
Thanks are also due to the South Australian
Museum, for the courtesy of Wolfgang Zeidler,
curator of Crustacea, and to the National Mu-
seum of Victoria, for the loan of type material; to
Dr. Phillip Gibbs of the New South Wales State
Fisheries Department for material from the Es-
tuarine Benthic Survey; and to Dr. Deborah Dex-
ter, Visiting Scientist at the Department of Zo-
ology, University of Sydney, 1980-1981, who very
kindly loaned to us some most useful material
from her New South Wales beach sands collec-
tion.
In the laboratory in Melbourne, Danuta Kar-
pow's technical assistance has been much appre-
ciated.
Dr. Alistair Gilmour, formerly Officer-in-
Charge of the Ministry for Conservation in Vic-
toria's Marine Studies Group and Dr. Leon Col-
lett, also of that Group, have supported this study
throughout. We are most deeply indebted to them
and to Dr. Barry Wilson, Director of the National
Museum of Victoria, who enabled the final year's
work in Melbourne to be carried out at that
institution.
We thank Dr. Thomas E. Bowman and Dr.
A.A. Fincham for suggestions to improve the
manuscript. We also are indebted to Jack Kory-
towski, our editor, for the many improvements
that he made to our work.
METHODS.?These fossorial taxa are treated by
the same methods and naming of taxa employed
by Barnard and Drummond (1978) for the Phox-
ocephalidae, with the following emphasis or dis-
tinction.
Spine and setal formulas read from proximal
to distal except in the case of article 3 of the
mandibular palp and of article 5 of pereopods 3
and 4 where they progress round the margin of
the article from anterior proximal to posterior
proximal.
Haustorioids are among the most setose of
amphipods and the setae are of many kinds. Some
are broad, short, and stiff and are called "spines,"
but other short blunt setae appear flexible or
bent; some are naked, blunt, and short and often
are inserted into second antennae by large basal
bulbs; other short setae are distally hairy or pu-
bescent and called "penicillate"; still others are
more evenly and sparsely hairy and called plu-
mose and these often are highly elongate and
flexible. We had difficulty, especially on article 1
of antenna 1, distinguishing between spines and
setae because the main row often has elements
too short and stout to be considered setae but too
bent or slightly flexible to be called spines; we
often use word combinations such as "spine-se-
tae" or "fine spines" to catalog the infinite degree
between extremes. Regardless of the appearance
of illustrations, the text description of spines is to
be accepted. We use the following code of abbre-
viations for setal formulas: E = long seta, often
plumose; e = short seta; P = penicillate seta; p
= penicillate setule; S = large spine; s = small
spine. A setule is a very small seta, especially the
small elements found on the telson.
NUMBER 360
"Glassy" spines are rodlike, stiff, untapering or
weakly tapering, often bifid spines attached to
the posterior margin of article 4 on antenna 2 in
Zobrachoidae and Urohaustoriidae; they do not
occur in the facial armament and so are excluded
from any formulas; a code letter is therefore not
used for them.
Marginal spines on palp article 3 of the man-
dible are cited in setal formulae in 4 groups:
dorsal (outer), apical, ventral (inner), any subap-
ical facial spines.
Thoracic appendages (legs 1-7) are termed
"gnathopods 1-2" followed by "pereopods 3-7";
other specialists may call these all "pereopods 1-
7" and not distinguish gnathopods 1-2 as sepa-
rate entities. Setation patterns on article 2 of legs
1-4 are so complex and variable that they have
been omitted.
Spine and setal formulas on pereopods 3-4 are
cited as in the following example: "5, 4, 2-1-1, 7
+ 6 + 1," in which "5" and "4" refer to large
apicoanterior setae on the outer face of articles 4
and 5; the formula "2-1-1" refers to posterior
spination on article 5, from distal to proximal,
the distal spine position being counted as paired;
"7 + 6 + 1" refers to the spines on article 6, with
7 lateral, 6 medial, and 1 intercalated between
the other rows.
Pleopodal formulas: peduncular formulas are
presented generally as "2 and 1, 2 and 0, 2 and
1" referring to the presence of 2 coupling hooks
on pleopods 1-3 plus an additional straight spine
on pleopods 1 and 3. Segmental formulas, cited
as 20-15, 18-12, 20-13, refer to segments on the
outer and inner ramus of the 3 pairs of pleopods.
Basal setal formulas follow the same pattern cit-
ing the number of setae on the basal articles of
the rami; for example, the formula "7-2-5-1"
means 7 lateral setae on article 1 of the outer
ramus, 2 medial setae on the same article, 5
lateral setae on article 1 of the inner ramus and
1 medial seta on the same article; formulas for
pleopods 2 and 3 then follow.
The armament formulas on the lateral margin
of article 1 on the outer ramus of uropod 1 and
on the antennae, proceed from proximal to distal.
Telsonic setal formulas cite one side of the
telson from base to apex, for example, "2P, 1, P,
2" refers to a pair of penicillate setules on the
basolateral margin followed by a marginal seta,
a penicillate seta and two apical setae; the total
setae of the telson are normally double the for-
mula. These armaments are usually very small
but use of small letters is reserved for elements
only in context and not in comparison to other
appendages.
Cuticular structures such as bulbar setules and
faint patterns of scales have been of so little value
in this study that they are largely ignored in the
descriptions but occasionally are illustrated.
Unknown characters are enclosed in bracket.
ORGANIZATION OF ILLUSTRATIONS.?Each figure
has one master specimen on which no subscript
lower case labels are placed to the left of the
capital letters denoting the kind of appendage.
Subsidiary specimens are marked with such labels
which are described in each legend.
DEFINITION OF TERMS.?Several concepts re-
quire explanation to enable use of terminology
that saves space and provides instant recognition
of a morphological state.
Antenna 1 of Haustorius Form: Peduncle
stout, short, articles 1-3 progressively shorter,
article 3 never elongate, flagella elongate (Figure
2: -41).
Antenna 1 of Urothoe Form: Peduncle slen-
der, one or more articles elongate, article 3 usually
elongate, flagella short (Figure 55: Al).
Antenna 2 of Haustorius Form: Article 4 of
peduncle broadly expanded, article 5 signifi-
cantly shorter and narrower than article 4 (Figure
2: A2).
Antenna 2 of Urothoe Form (and also of pon-
toporeia form): Article 4 of peduncle normally
rectilinear, article 5 not significantly shorter or
narrower than article 4 (Figure 55: A2).
Pereopod 5 of Haustorius Form: Articles 4
and 5 expanded but article 4 dominant (Figure
4: P5). Alternative is "pereopod 5 not of hausto-
rius form."
Uropod 3 of Ordinary Haustorioid-Phoxoce-
phalid Kind: Peduncle short and broad from
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
dorsal view, outer ramus longer than peduncle,
with 2 articles, first article dominant and usually
laterally armed, article 2 with apical armaments;
inner ramus variable, in female often very short
and poorly armed, but in male as long as outer
ramus and strongly setose; outer ramus in male
also strongly setose (Figure 17: R3).
Owing to the large size of the posterior pereo-
pods in many haustorioids, a small increase in
body length results in a great increase in massive-
ness of the specimens; hence, our attribution of
the terms "superadult" or "senile" may appear
strange in terms of the small increment in body
size of specimens so designated.
Names of new taxa are taken from aboriginal
words as follows: bannius, summit; buchalius, up-
right; Bumeralius, spirit; cangellus, good; Condukius,
spirit; Dirimus, spirit; garbaius, thin forehead; Ghee-
gerus, west wind; gunni, digging tool; karkan, spear;
merkamus, strange; Narumus, spirit; parnggius, deep
water; pentmus, withered; perkeus, steep; Prantinus,
jump with fright or wonder; pulcus, toed; talanggi,
linguiform; tallerkus, thistle; tarlitus, small; Tottun-
gus, evanescent; Tuldarus, astral; tungus, tarsal;
wingaro, oculate; yurrus, cloudy. All of the above
names are masculine.
TEXT ABBREVIATIONS
AM The Australian Museum, Sydney
CPBS Crib Point Benthic Survey, Western Port (sta-
tions in Barnard and Drummond, 1978)
EBS New South Wales State Fisheries Estuarine
Benthic Survey (Botany Bay, Jervis Bay, Lake
Macquarie, Merimbula)
HDWBS Hunter District Water Board Survey
NMV National Museum of Victoria, Melbourne
PPBES Port Phillip Bay Environmental Study (stations
in Barnard and Drummond, 1978)
QM Queensland Museum
QUBS University of Queensland Benthic Survey
SAM South Australian Museum, Adelaide
SBS Australian Museum Shelf Benthic Survey
WPBES Westernport Bay Environmental Study
F I G U R E A B B R E V I A T I O N S
(used in specimen drawings)
.1 antenna C head
B prebuccal from lateral /) dactyl of pereopod
/?;
F
C
11
I
J
K
L
M
N
<)
P
0.
R
S
T
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
maxilliped
telson
U
V
w
X
Y
Z
upper lip from anterior
palp
pleon (pleonites 1 3
bearing epimera;
pleonites 4-6, also re-
ferred to as urosomites
1-3, comprising uro-
some; often shown
with attached struc-
tures: telson and uro-
pods 1-3)
maxilla
see legend under illustra-
tion
see legend under illustra-
tion
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 ' V ) indicate the
following:
a alternate view
d dorsal
e broken
/flat
i medial
j unenlarged
/ lateral
o opposite
s setae removed
t right
v ventral
y enlarged
Superfamily HAUSTORIOIDEA Stebbing, 1906
Furnished with various fossorial (digging and
mud roiling) adaptations in form of extremely
setose antennae and appendages. Pereopod 5 of
special haustorius form (if not, rostrum cylindri-
cal), article 4 dominant, with strong facial spi-
nation on articles 4-5 (or progressively losing
spination in derived forms otherwise clearly con-
nected incrementally to plesiomorphic forms).
Uropod 3 of gammarid form (or progressively
modified, especially Japara Barnard and Drum-
mond, 1978), outer ramus dominant, 2-articulate,
if peduncle appearing elongate then rami
shortened. Accessory flagellum present, 2+ artic-
ulate. When entire, telson never longer than
broad and lamelliform; otherwise telson cleft. If
rostrum present then dorsoventrally flattened and
visorlike or thick, cylindrical and extended for-
NUMBER 360
ward without basal pediculation. Mandibular
palp 3-articulate.
If article 1 of flagellum on antenna 1 elongate
then gnathopods subchelate. Mouthpart field
quadrate (not conically grouped) from lateral
view, mouthparts not styliform. Mandibles with
incisor, otherwise not of pardaliscid form. If molar
weak then either of gnathopods 1-2 subchelate.
Outer plate of maxilliped well developed, its ar-
ticular base not elongate, inner plates if partially
fused, remaining broad, palp 3+ articulate. If
coxa 1 small, coxa 2 never as much as 4 times
longer or 10 times greater in area, coxa 2 never
larger than coxae 3 or 4. Gnathopods never car-
pochelate, together not of liljeborgiid form, article
3 not elongate. If partially fused together, uro-
somites not of tall form found in dexaminids or
ampeliscids.
CLASSIFICATION IN HAUSTORIOIDEA
In restricted terms and in modern context,
haustorioid amphipods were first segregated at
family level by Dana (1853), who instituted a
family Pontoporeiidae without definition but
with the inclusion of the following genera (spell-
ings here corrected): Pontoporeia, Pnscillina, Argissa,
Bathyporeia, Urothoe, Phoxocephalus, Harpinia, and
Lepidactylis. Stebbing (1906) removed Argissa to
the Argissidae and Phoxocephalus and Harpinia to
the Phoxocephalidae and used the name of the
oldest genus, Haustorius, as the family root, thus
constituting Haustoriidae with Bathyporeia, Pla-
tyischnopus, Haustorius, Cardenio, Pnscillina, Pontopor-
eia, Urothoe, and Urothoides. In recent times Cardenio
has been removed to the Synopiidae (Tironidae)
and Platyischnopus has been made the type of a
new family, Platyischnopidae (see Barnard and
Drummond, 1979). We do not believe Pontoporeia
and Haustorius are confamilial and we therefore
remove Pontoporeia and its cohorts to Gammaro-
idea.
Bousfield (1965) reinstated Pontoporeiidae as
a subfamily, Pontoporeiinae, to distinguish its
components, Pontoporeia, Amphiporeia, Bathyporeia,
Pnscillina, Haustoriopsis, and Urothoe, from the
Haustoriinae, which included Haustorius, Eohaus-
tonus, Protohaus(onus, Parahaustorius, Neohaustonus,
Pseudohaustonus, and Acanthohaustonus. The genera
Cardenio, Urohaustorius, Urothoides, Platyischnopus,
and Phoxocephalopsis were left unordered; Hausto-
riopsis has since been removed to the Dexaminidae
(see J. L. Barnard, 1972); and, as mentioned
above, Cardenio and Platyischnopus have been re-
aligned.
There is little doubt that the Haustoriinae,
sensu stricto, represent a very distinct and tightly
defined cluster of genera from the Atlantic
shorelines. They share such unusual characters as
absence of a maxillipedal dactyl and absence of
mandibular lobes on the lower lip. All have very
broad bodies and are immensely setose, with
powerful fossorial mechanisms. There is great
uniformity in thoracic appendages, pereopods 6
and 7 having a very stable, almost unique mor-
phology (see Bousfield, 1965). They exhibit a
uniform combination of other characters, some of
which do occur in non-haustoriin taxa, but not in
the following combination: short, broad head of
non-gammarus form; haustorius-type antennae;
reduced accessory flagellum; medial sides of max-
illae and maxillipeds strongly setose (maxilla 1
and maxillipeds often with basal baler lobes);
biarticulate palp of maxilla 1, enlarged outer
plate of maxilla 2; short, blunt mandibular inci-
sors; simple gnathopod 1; parachelate gnathopod
2; deeply lobate article 5 of pereopods 3 and 4;
haustorius-type pereopod 5 dominated by pereo-
pods 6 and 7; especially expanded article 4 of
pereopod 6; minute (or absence altogether of)
dactyls 3-7; short, broad pleopodal peduncles;
spinose, clavate or styliform rami on uropod 1;
linguiform setose rami on uropod 2; heavy ar-
mament at the joint of articles 1 and 2 of uropod
3; and setose apex of article 2. In addition epi-
meron 1 is well developed and epimeron 3 the
dominant side-plate. Brood plates, in contrast to
the narrow form of most antiboreal taxa. are
slightly expanded. Haustoriin telsons are more or
less similar, with a propensity for strongly gaping
but shallow apical cleavage and often a narrow
base. There is a tendencv to ventral flexion of the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
urosome, a condition not observed in pontopor-
eiins or in any antiboreal genera.
If one confines the pontoporeiin group to Pon-
toporeia, Amphiporeia, and Bathyporeia, a great con-
trast to haustoriins is evident. The head is of the
normal gammarus-form typical of basic gammar-
ids; the body is of the slender gammarus-form,
with the urosome, though slightly reduced in size,
unflexed; antenna 1 is of the gammarus- or
urothoe-form; the lower lip bears mandibular
lobes; pereopod 5 is not of the haustorius form;
the pleopods have elongate peduncles; uropod 2
has styliform or lanceolate rami bearing spines;
and the form of pereopods 5-7 and gnathopods
is variable. Other characters in common include
very short accessory flagellum, medial setation on
maxillae and maxillipeds and more or less ordi-
nary gammarid telson. Pnscillina can be admitted
to the Pontoporeiinae on the basis of maxillae,
lower lip, pereopod 5, pleopods, uropods, brood
plates (broad), and telson (though poorly cleft);
but the accessory flagellum is longer than in the
other genera and the head bears a sharp ventral
cheek as in Urothoe.
Haustoriinae and all other haustorioid taxa,
including Urothoe, Urothoides, Carangolia, Cumcus,
Phoxocephalopsis, Zobracho, Bumeralins, Gheegerus,
Prantinus, Dinmus, Narumus, Urohaustorius, Tuldarus,
and Tottungus differ from Pontoporeiinae in the
haustorius form of pereopod 5, the short pedun-
cles of the pleopods, and the presence of thick
posterior spines on article 5 of pereopods 3-4, but
in the haustoriins article 5 is broadly expanded
and the spines occur in 2 distinct rows. Except
for Haustoriinae and Phoxocephalopsis, all genera
have these spines in a distinct pattern: a pair of
distal spines and one or more posterior marginal
spines.
If Phoxocephalopsis is held in abeyance, the non-
haustoriin genera are divisible into two large
groups, the Urothoe group with urothoe antenna
2, and the Urohaustorius group with haustorius
antenna 2. The Urothoe group is further charac-
terized by the large ventral cheek on the head
and the Urohaustorius group by the presence of a
special set of numerous glassy, untapering setal-
spines occurring ventrally and ventro-facially on
article 4 of antenna 2. When rami of uropods 1,
2 are present and well developed they are lingui-
form and setose in the Urohaustorius group and
styliform-lanceolate and spinose in the Urothoe
group. Most members of the Urohaustorius group
have the inner rami of the pleopods significantly
shorter than the outer. In the genus Urothoe (but
unknown for others of its group) the rami are
subequal. All members of the Urohaustorius group
are also characterized by the dominance of epi-
meron 2.
In these terms the Urothoe group includes Uro-
thoides, Carangolia, and Cumcus, and the Urohausto-
rius group includes Zobracho, Bumeralius, Narumus,
Prantinus, Gheegerus, Dinmus, Tottungus, and Tulda-
rus. Within the Urohaustorius group a subdivision
based on simple and subchelate gnathopod 1 and
discreteness of epimeron 1 can be made, with the
subchelate gnathopod 1 and discrete epimeron I
being found in Zobracho, Bumeralius, and Prantinus.
In Prantinus, however, epimeron 1 is least discrete
and approaches the condition found in Urohaus-
torius and its allies.
Antenna 1 of all members of the Urothoe group
is of the urothoe form, and of all members except
Prantinus of the Urohaustorius group is of the haus-
torius form. Prantinus remains more strongly as-
sociated with the Urohaustorius group than with
the Urothoe group on the basis of antenna 2 and
the rami of uropods 1-2.
Phoxocephalopsis is enigmatic. It combines the
Haustorius antennae 1-2, maxillary palp, pereo-
pod 5, and uropods 1 and 2 with the Urohaustorius
kind of maxillipeds and other mouthparts (except
palp of maxilla 1), pereopods 3-4, pereopods 6-7,
epimera, and telson. It cannot be grouped with
Urothoe because of antennae 1-2 and uropod 2.
The head check is moderate and the special glassy
spines of antenna 2 are absent. It would appear
to be a plesiomorphic form standing ancestral
both to the Urohaustorius group and the Hausto-
riinae. It deserves familial rank.
All members of the greater Urothoe-Urohaustonus
group occur in the southern hemisphere except
for several species of Urothoe that have penetrated
NUMBER 360
the northern hemisphere and generally live in its
cooler waters or in the tropical waters of southern
and eastern Asia. All pontoporeiins and hausto-
riins are centered on the north Atlantic Ocean.
Pontoporeiins and haustoriins bear widened
brood plates whereas all known urothoin-uro-
haustoriin taxa have narrow brood plates.
EVOLUTIONARY PATTERN IN HAUSTORIOIDEA
FIGURE 1
The details given in the classificatory section
suggest the broad outlines of an evolutionary
pattern. The various groups are raised to family
level.
The pontoporeiids would appear to be the most
primitive of the group but this is not to suggest
that they are ancestral to the others. The differ-
ences are so strong that one may not conclude
that any evolutionary flow has occurred between
pontoporeiids and other haustorioid groups. Pon-
toporeiids are very close to the Pontocaspian gam-
maroids (see discussion in Barnard and Drum-
mond, 1978) but differ in the enfeeblement of the
gnathopods and the somewhat enlarged mandib-
ular molars with relatively weaker triturative
states and the loss of coxal gill 7. The pontopor-
eiids are so close to the Pontocaspian gammaroids
that they probably do not deserve more than
minor classificatory disjunction from the
"Pontogammaridae" (family considered invalid;
see Barnard and Barnard, in press). Whatever
intergradation could be suggested to occur be-
tween pontoporeiids and the remaining hausto-
rioids can be seen in Priscillina, on which antenna
2 is weakly of the haustorius form, with a few
facial spines and reduced article 5 and on which
a pointed ventral cephalic cheek occurs similar
to, but possibly not homologous with, that found
in Urothoe. Priscillina also resembles the urothoids
in the blunted mandibular incisors. In contrast to
Priscillina, antenna 1 and pereopods 5-7 are very
distinctive in urothoids. Pleopodal peduncles of
Priscillina (new observation) are somewhat inter-
mediate in their elongation between those of Pon-
toporeia and Urothoe.
The urothoids, comprising Urothoe, Urothoides,
Carangolia, and Cumcus (and probably more gen-
era to be separated out of Urothoe), differ from
pontoporeiids in the special form of antenna 1,
on which one or more articles are elongate, espe-
cially article 3, and on which the flagella are very
short; the weak medial setation of maxilla 1; and
the special haustorius form of pereopod 5, on
which articles 4-5 are broadened but article 4 is
dominant. In the most primitive urothoids the
lateral faces of these articles are densely spinose
but in some of the more apomorphic species, such
as Urothoe elegans, many of the spines are lost and
article 4 scarcely dominates article 5. The biarti-
culate palp of maxilla 1, typical of pontoporeiids,
is retained. Articles 5 and 6 of pereopods 3-4
exhibit a pattern of spination not found in pon-
toporeiids. Fully developed facial armaments, in
ranks, are developed on article 4 of antenna 2.
Like Bathyporeia of the Pontoporeiidae, many uro-
thoids have strong sexual dimorphism in antenna
2, the flagellum of the male becoming elongate
and calceoli being developed on the peduncle and
the flagellum. The head of urothoids is recogniz-
able because of the ventral protrusion of a sharp
cheek similar to that of Priscillina.
A more or less orderly progression in the devel-
opment of the urothoid kind of pereopods 3-4
through the Phoxocephalidae may be seen; the
primitive phoxocephalid has an elongate article
6 and poorly spined article 5, whereas such ad-
vanced taxa as Yammacoona and Kotla (Barnard
and Drummond, 1978) have the strongly spined
article 5, though article 6, with its similar spina-
tion pattern, never becomes as broad as in uro-
thoids. Those genera also have a severely reduced
rostrum, dorsoventrally extended head, and uro-
thoid antenna 1; and retain the biarticulate palps
of maxilla 1. One might visualize an ancestor of
Kotla as a simultaneous ancestor of Urothoides, in
which case gnathopodal enfeeblement and slight
lessening of distinctions between pereopods 6 and
7 would have to ensue in the urothoid group.
Because it would be difficult to derive other
haustorioids from the same point as Kotla, the
support for manifold derivation (polyphyly) of
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
GAMMARIDAE
PONTOPOREIIDAE
UROTHOIDAE
PHOXOCEPHAUDAE-*
PLATYISCHNOPIDAE
CONDUKIIDAEPHOXOCEPHALOPSIDAE
ZOBRACHOIDAE
UROHAUSTORIIDAEHAUSTORIIDAE
Heac
Antenna 2
M a x i l l o 1 Pe r ecpod 5
Pereopod 3
Lower
Uropod 1 Uropod 2
G l a s s y S p i n e s
M a n d i b u - o r P c i p
P i e o p o d
OXP
C i o t h e s p i r
S p . n e
FIGURE 1.?Evolutionary- deployment in various families of the hypothetical haustorioid group.
NUMBER 360
the various groups of haustorioids is amplified.
The zobrachoid-urohaustoriid line would have a
better starting point near the primitive end of the
phoxocephalid scale, but again article 6 of per-
eopods 3-4 would develop convergently.
All remaining haustorioids are characterized
by the haustorius antenna 2, in which article 4 is
expanded and article 5 is greatly inferior. All
except Prantinus are also characterized by the
haustorius antenna 1, which resembles that of
pontoporeiids, but in which the accessory flagel-
lum is enlarged. At this point one must dispose of
the Phoxocephalidae, because the primitive mem-
bers and the majority of Phoxocephalidae are
characterized by the combination of haustorius
antennae 1-2, and pereopod 5. Like Bathyporeia
and Urothoe many of the taxa have the elongate,
sexually dimorphic male antenna 2. Unlike pon-
toporeiids, urothoids, and other haustorioids, sev-
eral phoxocephalids retain enlarged gnathopod
2, a possible mark of their ancestry in gammarids.
The short wrist of this phoxocephalid gnathopod
2 is unlike that of gammarids, however, and may
indicate a secondary evolutionary development.
Though most phoxocephalids have enfeebled
gnathopods and small, non-triturative mandibu-
lar molars, several genera retain much better-
developed triturative mandibular molars than do
any haustorioids outside of the pontoporeiids.
This exceptional molar is much more acetabular
than any found in gammarids. Urothoids and
phoxocephalids share much in common, includ-
ing a slight propensity in Urothoe for the devel-
opment of a visorlike rostrum, the characteristic
plesiomorph of phoxocephalids. Nevertheless,
Barnard and Drummond (1978), on the basis of
the more plesiomorphic gnathopods and mandi-
bles of phoxocephalids, prefered not to derive
phoxocephalids from urothoids. There is a possi-
bility that these had a common ancestor, but no
idea of straight-line derivation can be enter-
tained. Urothoids retain the unusual palp of max-
illa 1 found in pontoporeiids and in the sensu
stricto Haustoriidae, in which, quite unlike most
amphipods, and especially unlike gammarids and
phoxocephalids, article 1 is elongate. Occasional
phoxocephalids possess a urothoid antenna 1,
though rarely with article 3 as elongate as in
urothoids.
The next most apomorphic group would ap-
pear to be the provisional "urohaustoriids," com-
prising Zobracho, Bumeralms, Prantinus, Gheegerus,
Narumus, Urohaustorius, Tuldarus, Dinmus, and Tot-
tungus. Actually this group stands opposite the
Haustoriidae in apomorphy. Its uropods 1-2 and
maxilla 1 are much more specialized than those
of the haustoriids, whereas in haustoriids pereo-
pods 6-7, uropod 3, and mouthparts other than
maxilla 1, show greater specialization than in the
Urohaustoriidae. Neither group could be derived
from the other without severe evolutionary rever-
sals in various characters, though they can be
visualized to have a common ancestor.
All urohaustoriids are characterized by the lin-
guiform setose rami of uropods 1-2; this kind of
ramus in Haustoriidae occurs only on uropod 2.
Urohaustoriids are also characterized by the uni-
articulate palp of maxilla 1, the clustering of the
glassy spines on article 4 of antenna 2, the domi-
nance of epimeron 2, the presence of large or
easily visible elements forming an armament row
on the outer face of article 1 on antenna 1 (only
remnants occurring in Haustoriidae), the specific
form of pereopods 6-7, (articles 4-5 poorly ex-
panded and article 4 on pereopod 6 not hugely
dominant), the presence of clothespin spines on
the inner rami of the pleopods, and by the absence
(therefore normalcy) of various other characters
to be mentioned specifically for Haustoriidae be-
low. Some urohaustoriids also bear the clothespin
spines on the pleopods; these may be a primordial
attribute lost by Haustoriidae.
Urohaustoriids are divisible into 2 groups:
those without palm on gnathopod 1, which are
typical of I'rohaustorius and its allies: and those
with subchelate gnathopod 1. like Zobracho. Bum-
eralms, and Prantinus. These 3 are segregated into
the family Zobrachoidae but otherwise they stand
close to Urohaustoriidae. One assumes that sub-
chelate gnathopod 1 is the plesiomorphic state
and that urohaustoriids arc therefore highly apo-
morphic; but Prantinus is so divergent and apo-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
morphic in other attributes that it may have to
be split away into a group of its own. Antenna 1
of Prantinus is of the urothoe form but in most
other features Prantinus resembles Zobracho. Pran-
tinus differs from any urothoid in the incisor and
molar of the mandible, the dominance of epime-
ron 2, and in characters of the inner plate and
palp of maxilla 1.
Because Cumcus is so divergent, with its unusual
uropods 1-2, composed only of setose peduncles,
the urothoids cannot be limited by the character
of styliform rami on uropod 1. One infers that an
hypothetical evolutionary step occurs between
styliform spinose rami and the loss of rami (with
attendant linguiform-setose peduncles). This
might be represented by a stage similar to that
seen in Prantinus, where the linguiform rami have
lost all but 1 apical seta.
Haustoriids are probably the most specialized
of the haustorioids because of numerous oddities
in the mouthparts rarely encountered in other
amphipods. Although they retain a biarticulate
palp on maxilla 1 and strongly setose medial
edges of maxillae and maxillipeds, many species
develop baler lobes basally on maxilla 1 and the
maxillipeds. Palp article 4 of the maxilliped is
absent and article 3 is strongly geniculate. The
outer plate of maxilla 2 is often greatly enlarged.
The lower lip lacks mandibular lobes. The man-
dibular incisors are blunted and palp article 3 of
the mandible retains at least 3 kinds of setae in 2,
3, or 4 groups (interpretive). The apical setae on
palp article 3 are not awned as in most urohaus-
toriids; most are simple or weakly hooked api-
cally, though the inner basal members of the
apical group often are bilaterally serrate and
apically sheathed (new observations) and the in-
ner basal margin bears a row of setae that are
strongly serrate bilaterally. This kind of seta is
not found in the urohaustoriids, where the setae
of palp article 3 are mixed between awned and
simple, and no special inner basal row is present.
Haustoriids further differ from urohaustoriids
in the loss of all but remnants (minute penicillate
setules) in the armament row on article 1 of
antenna I: the shortened accessory flagellum (also
weakly occuring in the urohaustoriid, Tottungus);
the evenly spread, tandemly spaced, simple glassy
spines on article 4 of antenna 2 (bifid and clus-
tered in urohaustoriids); the absence of button
combs (Figure 21: QGU, arrow) on the gnatho-
pods; the development of a large posterior lobe
on article 5 of pereopods 3-4; the extreme vesti-
giality of dactyls on all of pereopods 3-7 (repre-
sented only by a small button and seta appearing
as a bulbar setule, new observation); the full
flattening of article 6 on pereopods 3-4, with the
resultant wide spacing in the spine rows; the
broader brood plates; the special character of
pereopods 6-7; the longer inner rami of the pleo-
pods and the absence of clothespin spines on these
inner rami; the lack of dominance by epimeron
2; the clavate or rodlike rami of uropod 1 (not
linguiform); and the rodlike rami of uropod 3,
with multispinosetation apically on both rami
and on article 1 of the outer ramus.
To some extent the general form of the telson
in haustoriids is distinct from that in urohausto-
riids, but the telson in the latter is so variable that
the differences are subjective.
Once the distinctions between haustoriids and
urohaustoriids, as noted above, can be visualized,
the primitive position of Phoxocephalopsis, from
antiboreal South America, is apparent (Figure 1).
Several of its attributes were not described pre-
cisely but the description and figures of Schellen-
berg (1931) and a new analysis (page 15) are
adequate to distingish the genus from urohaus-
toriids, urothoids, and haustoriids. Although the
palp of maxilla 1 is biarticulate, and uropods 1-
2 are much like haustoriids, Phoxocephalopsis is far
more similar to urohaustoriids than to hausto-
riids. Epimeron 2 is weakly dominant, the rami
of uropod 3 are not rodlike, article 5 of pereopods
3-4 is not grossly expanded as in haustoriids, all
pereopodal dactyls are well developed, pereopods
6-7 resemble those of urohaustoriids more closely
than of haustoriids, and the maxillae and maxil-
lipeds are almost naked medially. The accessory
flagellum, however, is somewhat short. The fine
structure of the mandibles mixes urohaustoriid
incisors, laciniae mobiles, and rakers, with haus-
toriid molarial texture and condition of palpar
NUMBER 360 11
spines. The lower lip bears mandibular lobes. The
gnathopods resemble those of urohaustoriids.
Phoxocephalopsis differs from urothoids in the
haustorius form of the antennae but the cephalic
cheek is weak. Schellenberg appropriately named
the genus for its resemblance to phoxocephalids
in rostrum, but pereopod 7 is not of the phoxo-
cephalid form, epimeron 2 is dominant, and,
unlike phoxocephalids, palp article 2 of the max-
illipeds is expanded.
If a rule of evolutionary irreversibility is
adopted then none of the known haustorioids
represents a perfect intergrade among the several
groups, Pontoporeiidae, Phoxocephalidae, Uro-
thoidae, Phoxocephalopsidae, Urohaustoriidae,
Zobrachoidae, and Haustoriidae. These inter-
grades, lying in the crotches of an evolutionary
tree, must be hypothesized. Examples of necessary
reversal in evolutionary trends, as a step towards
Urohaustoriidae, may be cited as: the need for
reversal in urothoid antenna 1 to the pontopor-
eiid-haustoriid condition; the reversal of domi-
nance in epimeron 2 as a step between Phoxoce-
phalopsidae and Haustoriidae; the reversal in
linguiform rami of uropods 1-2 towards the styl-
iform spinose condition as a step between Uro-
haustoriidae and Urothoidae; the redevelopment
of mandibular lobes on the lower lip; and the
creation of a fourth palp article on the maxillipeds
of Haustoriidae as a step towards Urohaustori-
idae.
The minimum morphologic state of the most
primitive haustorioid (discounting Pontoporei-
idae) would require styliform rami of uropods 1-
2, subchelate gnathopod 1, antenna 1 of the
haustorius form, antenna 2 of the urothoid-gam-
marid form, maxilla 1 with biarticulate palp,
maxillipeds with dactyls, and pereopod 5 of the
haustorius form. If Phoxocephalidae are to be
included in the scheme and if their so-called
plesiomorphic states in gnathopods and molar are
to be accepted as truly primitive and not second-
arily derived, then a primitive member would
require fully triturative molars and enlarged
gnathopod 2. One would then assume that the
special pereopod 7 and rostrum of Phoxocephal-
idae and Zobracho are secondary developments, as
are the cheeks of urothoids, the linguiform uro-
podal rami of urohaustoriids, and the odd first
antennae of urothoids and Prantinus. No living
pontoporeiid fits this pattern.
Platyischnopids, like harpiniins within the
Phoxocephalidae, have a modified haustorius
pereopod 5. Articles 2, 4, and 5 are poorly ex-
panded, and articles 4-5 poorly spined facially.
In other respects (the special form of the incisors,
resembling apomorphic phoxocephalids, the
shape of coxa 4, maxilla 1 generally, epimera,
uropods, and to some degree the chelate gnatho-
pods), platyischnopids bear close resemblance to
members of the Phoxocephalinae, one of the most
advanced groups of the Phoxocephalidae. The
definition of Platyischnopidae could be simplified
to the statement: "Phoxocephalids with cylindri-
cal rostra." The special form of the mandibular
incisor cannot be accorded absolute homology to
that of phoxocephalids because the same kind of
incisor crops up in Oedicerotidae (for example,
Synchelidium) and may be polyphyletic.
Condukiids disturb the tight definition of
Haustorioidea, because, like Pontoporeiidae (now
assigned to Gammaroidea), pereopod 5 is not of
the haustorius form, rather it is a replica of the
combined formation found on pereopods 6-7 in
haustoriids, with article 5 greatly dominant and
facial spines severely reduced.
A discussion of hypothetical states and trends,
reversible or not, could go on endlessly. Suffice it
to say that the final absurdity is the lack of proof
that the haustorioid pereopod 5 is not a replica-
tory character of convergence, with several
origins. Until this can be determined there is little
point in attempting to trace exact evolutionary
webs by supposition of missing links.
In summary, we have cast doubt on the mon-
ophyletic state of pontoporeiids and other haus-
torioids and must suggest that pontoporeiids be
closely allied to the "Pontogammaridae" (includ-
ing "Compactogammarinae"). while provision-
ally the Haustorioidea may encompass Urothoi-
dae, Phoxocephalopsidae, Urohaustoriidae, Zo-
brachoidae, Phoxocephalidae, Platyischnopidae.
and Condukiidae. The latter two groups may
require their own superfamily.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Key to Families of Haustorioidea
1. Rostrum cylindrical 2
Rostrum not cylindrical or absent 3
2. Article 5 of pereopods 3-4 strongly produced posteriorly, peduncles of
pleopods wider than long or as wide as long. Condukiidae, new family
Article 5 of pereopods 3-4 unproduced posteriorly, peduncles of pleopods
longer than wide Platyischnopidae
3. Head with sharp ventral cheek 4
Head lacking sharp ventral cheek 5
4. Antenna 1 with article 3 at least half as long as article 1, mandibular
molars large and pubescent, incisors broad and almost toothed, pedun-
cles of pleopods not longer than wide Urothoidae (part)
Antenna 1 with article 3 very short, mandibular molars triturative or not,
but not large nor pubescent, incisors broad but 3-4 toothed, peduncles
of pleopods longer than wide Phoxocephalidae (part)
5. Pereopod 7 of characteristic phoxocephalid form
Phoxocephalidae (part)
Pereopod 7 not of phoxocephalid form 6
6. Rami of uropod 1 styliform and either naked, spinose, or rami absent;
palp of maxilla 1 biarticulate 7
Rami of uropod 1 linguiform and setose, palp of maxilla 1 uniarticulate
9
7. Rami of uropod 2 linguiform and setose, palp of maxilliped 3-articulate
Haustoriidae
Rami of uropod 2 rodlike and spinose or rami absent, palp of maxilliped
4-articulate 8
8. Antennae of haustoriid form, article 1 of antenna 1 short, of antenna 2
broadly expanded Phoxocephalopsidae
Antennae of urothoid form, article 1 of antenna 1 elongate, article 4 of
antenna 2 poorly expanded Urothoidae (part)
9. Gnathopod 1 subchelate Zobrachoidae , new family
Gnathopod 1 simple Urohaustoriidae, new family
PHOXOCEPHALIDAE
DIAGNOSIS.?Rostrum forming huge anterior
visor, not cylindrical, reduced in apomorphic gen-
era, cheek occasionally well developed. Antenna
1 of haustorius form, apomorphically of urothoe
form, accessory flagellum rarely elongate. An-
tenna 2 of modified urothoe form, article 5 often
very small but spines on face of article 4 organized
into ranks, ventral margin lacking glassy spines
of haustorius form, flagellum usually longer than
article 4 of peduncle, often greatly enlarged in
male, and antenna 2 then bearing calceoli. Pre-
buccal complex rarely massive, upper lip usually
dominant. Mandibles bearing short, broad inci-
sors usually with 3 teeth, 1 tooth widely disjunct,
laciniae mobiles present on both sides and unlike
each other, rakers numerous and usually serrate,
molar variable, fully triturative to vestigial and
poorly spinose, accessory chopper weak or absent,
palp 3-articulate, at most with several outer setae,
usually with very few, most other spines concen-
trated on inner apical level, spines not awned.
Lower lip with discrete inner lobes and strong
NUMBER 360 13
mandibular extensions. Maxilla 1 with 1-2 artic-
ulate palp, setosity of inner plate variable. Max-
illa 2 ordinary, inner plate poorly setose medially
and facially. Maxillipeds with unexpanded bases,
plates variable, outer spinose, palp 4-articulate,
article 2 not expanded in form of other hausto-
rioids, article 4 variable, unguiform to clavate.
No baler lobes on mouthparts.
Coxae 1-4 evenly integrated by stepped en-
largement from coxa 1 onwards, coxa 4 dominant,
coxa 1 large, coxa 3 not produced posteroven-
trally. Coxal gills on pereonites 2-7, brood plates
slender.
Gnathopods variable. Article 5 of pereopods 3-
4 slightly expanded, not deeply lobate, with or
without thick posterior spines, article 6 usually
narrow and elongate, rarely expanded; dactyls of
pereopods 3-7 usually well developed; pereopod
5 of haustorius form becoming modified apo-
morphically (losing breadth and spination); per-
eopods 6-7 not alike; pereopod 6 large, with
subovately expanded article 2; pereopod 7
smaller, with expanded, shieldlike article 2; apical
articles much shortened and stenopodous. No
pereopod with underslung articulation.
Pleopod 2 not inferior; peduncles of pleopods
longer than broad, coupling hooks of haustorius
form; inner rami subequal to outer, lacking
clothespin spines. Epimeron 1 well developed;
epimeron 3 dominant in size and setation.
Urosomites ordinary to slightly modified
(spiked or fused). Rami of uropods 1-2 styliform,
spinose; uropod 3 usually of ordinary haustorioid-
phoxocephalid kind, outer ramus dominant, usu-
ally 2-articulate, peduncle slightly elongate, rami
poorly setose apically; or uropod 3 variable, with
occasional modifications and reductions.
Telson long to short, cleft. Sexual dimorphism
often strong in antennae, urosome, uropod 3.
TYPE GENUS.?Phoxocephalus Stebbing.
COMPOSITION.?47 genera (Barnard and Drum-
mond, 1978).
PLATYISCHNOPIDAE
DIAGNOSIS.?Rostrum strong, cylindrical, with
subapical ventral retrorse process, cheek poorly
developed. Antenna 1 of haustorius form or
urothoe form (typical), flagella elongate. Antenna
2 of urothoe form, article 5 slightly to strongly
shorter and slightly narrower than article 4, latter
without major facial armament, ventral margin
with 1 kind of armament, either setae or glassy
spines, flagellum often greatly elongate in male
and antenna 2 then bearing calceoli. Prebuccal
complex massive, upper lip dominant, epistome
scarcely evident. Mandibles bearing short, broad,
phoxocephalid-like incisors with 3 main teeth or
stubby modifications thereof, 1 of these widely
disjunct; lacinae mobiles present on both sides,
alike or not, rakers moderately numerous and
serrate, molar of medium size, not triturative,
bearing spinules, lacking accessory chopper; palp
3-articulate, article 3 lacking outer setae, all
spines concentrated on apical bevel. Lower lip
with discrete inner lobes, mandibular extensions
of outer lobes well developed. Maxilla 1 with
uniarticulate palp, inner plate with fewer than 5
setae. Maxilla 2 ordinary or outer plate enlarged,
inner plate with poorly developed oblique facial
row of setae. Maxillipeds with unexpanded bases,
poorly enlarged plates, outer spinose; palp 4-ar-
ticulate, article 2 wide but not differentially ex-
panded, article 4 unguiform, with or without nail.
No baler lobes on any maxillae or maxillipeds.
Coxae variable but coxa 4 strongly dominant
and posterodorsally excavate, thus with poste-
riorly directed lobe, coxa 3 rectangular, or ven-
trally narrow, coxae 1-2 of various dimensions.
Coxal gills on segments 2-6, 3-7, or 2-7; brood
plates slender.
Gnathopods feeble or slightly enlarged, alike or
not, wrists more or less elongate, hands elongate
or not, strongly or weakly chelate. Article 5 of
pereopods 3-4 narrow, with sparse posterior spi-
nation; dactyls of pereopods 3-7 well developed
or pereopod 5 weakly of haustorius form, articles
2, 4, occasionally 5, expanded, articles 4-5 with
weakly developed facial spination; pereopods 6-
7 more or less alike but pereopod 7 larger, article
2 usually of modified phoxocephalid form, article
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
4 of pereopods 6-7 expanded; no pereopod with
underslung articulation.
Pleopod 2 not inferior. Peduncles of pleopods
much longer than wide, coupling hooks of haus-
torius kind; inner rami as long as outer (or very
nearly), bearing multiple clothespin spines with
small inner barbs. Epimeron 1 well developed;
epimeron 2 often dominant in size.
Urosomites ordinary. Rami of uropods 1-2 styl-
iform or rodlike, spinose; uropod 3 of phoxoce-
phalid form but inner ramus very small to mod-
erate in size, article 2 of outer ramus small to
greatly elongate, apices of rami poorly setose.
Telson variable, cleft to entire. Sexual dimor-
phism occuring on antenna 2 and uropod 3.
TYPE GENUS.?Platyischnopus Stebbing.
COMPOSITION.? Tomituka Barnard and Drum-
mond, Indischnopus Barnard and Drummond, Yur-
rokus Barnard and Drummond, Tittakunara Bar-
nard and Drummond; and undescribed Ameri-
can genera. (See Barnard and Drummond, 1979.)
RELATIONSHIP.?Platyischnopids have so many
characters in common with the subfamily Phox-
ocephalinae (Phoxocephalidae) that one might
derive them from an ancestor common to that
group. The structure of the mandibular incisors
and the pattern of spine distribution on the man-
dibular palp, plus the shape of coxa 4 suggest this
affinity. Article 2 of pereopod 7 has faint similar-
ity to that of phoxocephalids but the remainder
of the appendage is much stouter and more elon-
gate than in phoxocephalids. The unusual head
is a distinguishing mark.
See "Condukiidae" for further comments.
UROTHOIDAE
DIAGNOSIS.?Rostrum weak, head short, ven-
tral cheek strongly developed and projecting ven-
trally. Antenna 1 of urothoe form, articles 1-3
elongate, geniculate, flagella short. Antenna 2 of
urothoe form, article 4 scarcely expanded, with
spines in seriate ranks, ventral margin with 2
kinds of setae, plumes, and penicillates, ventral
glassy spines poorly developed or absent, article
5 slightly shorter and narrower than article 4,
flagellum variable, usually short in female but in
male often greatly elongate as in Bathyporeia or
phoxocephalids, and flagellum, plus part of pe-
duncle, often furnished with calceoli. Prebuccal
complex massive, epistome scarcely distinct, up-
per lip dominant. Mandibles bearing stubby,
poorly toothed incisors; laciniae mobiles present
on both sides and unlike each other, rakers poorly
developed to absent; molar large, minutely fuzzy,
almost non-triturative, lacking significant acces-
sory chopper; palp 3-articulate, article 3 with
outer setae, setae simple. Lower lip with discrete
inner lobes, mandibular extensions of outer lobes
well developed. Maxilla 1 with biarticulate palp,
inner plate with fewer than 5 setae. Maxilla 2
ordinary, inner plate with well-developed medial
to submarginal row of setae. Maxillipeds with
unexpanded bases, normally enlarged plates,
outer spinose; palp 4-articulate, article 2 ex-
panded, article 4 unguiform to clavate, usually
setose, apical nail usually poorly developed. No
baler lobes on any maxillae or maxillipeds.
Coxae variable, either of ordinary gammarid
form, or coxa 1 very small, or coxa 3 dominant
and/or bearing posteroventral lobe, plus other
variations. Coxal gills on segments 2-5 or 2-6;
brood plates slender.
Gnathopods feeble, grossly alike in proportions,
wrists elongate, but otherwise highly variable in
degrees of palmar development and chelateness,
article 3 short. Article 5 of pereopods 3-4 broad,
slightly expanded, not deeply lobate, with thick
posterior spines; dactyls of pereopods 3-5 well
developed, those of pereopods 6-7 variable but
usually well developed; pereopod 5 of haustorius
form, articles 2, 4, and 5 expanded, articles 4-5
with extensive facial rows of spines in primitive
members but declining in apomorphic taxa; per-
eopods 6-7 alike, or pereopod 7 developed in
phoxocephalid fashion, article 2 broadly ex-
panded and posteroventrally lobate, remainder
of articles thin, whole appendage somewhat
shortened in comparison to pereopod 6.
Pleopod 2 apparently not inferior; peduncles of
pleopods slightly to significantly wider than long,
coupling hooks paired on each pleopod; inner
NUMBER 360 15
rami usually inferior, usually not bearing
clothespin hooks (only several species in 2 genera
examined). Epimeron 1 strongly developed; epi-
meron 3 dominant in size and usually in setation.
Urosomites ordinary. Rami of uropods 1-2 styl-
iform and spinose or rami absent; peduncles usu-
ally only spinose, but 1 genus with linguiform
and setose peduncles lacking rami; uropod 3 of
ordinary haustorioid-phoxocephalid kind, outer
ramus dominant, 2-articulate, peduncle short,
flat, expanded; rami poorly setose apically.
Telson variable. Antenna 2 with strong sexual
dimorphism in several taxa.
TYPE GENUS.?Urothoe Dana.
COMPOSITION.? Urothoides Stebbing (see Bar-
nard and Drummond, 1979); Carangolia J.L. Bar-
nard; Cumcus Griffiths.
RELATIONSHIP.?Though 1 genus of Urothoi-
dae, Urothoides, bears the proper formula of per-
eopods 6-7 to be classified as a phoxocephalid,
the presence of a large ventrally extended ce-
phalic cheek shows its affinities to Urothoe, on
which pereopods 6-7 are not of the phoxoce-
phalid form. The mandibles of urothoids are
unlike those of phoxocephalids and though uro-
thoids have a small rostrum (smaller than Zo-
bracho), they never bear the fully developed ros-
trum of plesiomorphic phoxocephalids. One may
entertain the suggestion that urothoids could be
descended from phoxocephalids by alterations in
molar, rostrum, and cephalic cheek by intergrad-
ing them through Urothoides (which bears phoxo-
cephalid pereopods), but the reverse direction
would be highly unlikely. So-called primitive
phoxocephalids bear enlarged gnathopod 2 and
fully triturative molars; one would doubt that
such conditions could be descendent from uro-
thoids (Barnard and Drummond, 1978). The ma-
jority position of the haustorius antenna 1, not
only in the haustorioids but in the pontoporeiids
and the pontogammarids, suggests that it repre-
sents the plesiomorphic state, whereas the rarer
urothoid antenna 1, present in urothoids, Pran-
tinus, certain platyischnopids, and occasional
phoxocephalids, is apomorphic.
Key to Genera of Urothoidae
1. Pereopod 5 of phoxocephalid form Urothoides
Pereopod 5 not of phoxocephalid form 2
2. Uropods 1-2 lacking rami Cunicus
Uropods 1-2 bearing rami 3
3. Telson and uropod 3 stubby, very short Carangolia
Telson and uropod 3 of normal length Urothoe
PHOXOCEPHALOPSIDAE
DIAGNOSIS (new observations on South Ameri-
can material in italics).?Rostrum of moderate
size, broad; cheek weak. Antenna 1 of haustorius
form, article 1 short, main setal row strong, article 2
as long as article 1, article 3 shorter, primary
flagellum elongate, accessory flagellum short. An-
tenna 2 of haustorius form, article 4 expanded,
article 5 shorter and narrower than article 4, these
articles furnished with 1 or more longitudinal
rows of facial armaments, ventral margin with
long setae, and 2 other kinds but none glassy; flagellum
longer than article 4 of peduncle. Epistome and
upper lip distinct. Mandibles with stubby, sparsely but
distinctly toothed incisors, lacunae mobiles diverse, rakers
sparse and widely separate, molar of medium size, tritur-
ative, choppers weak; palp 3-articulate, article 3 with
urohaustonid setal distribution, setae hooked but un-
owned. Lower lip with mandibular lobes moderately well
developed. Maxilla 1 with biarticulate palp, inner
plate small and with one seta. Maxilla 2 ordinary,
inner plate with oblique facial row of setae. Max-
illipeds with unexpanded bases and no baler lobes, with
normally enlarged plates, outer spinose; palp 4-
articulate, article 2 expanded, article 4 clavate.
multisetose.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Coxae 1-4 apparently evenly integrated by
stepped enlargement from coxa 1 onwards, coxa
4 dominant, coxa 1 medium and slightly tapering,
coxae 3-4 not (type) or distinctly (other species)
produced posteroventrally. Coxal gills 2-6; brood-
plates slender.
Gnathopods feeble, grossly alike in proportions,
wrists elongate, but gnathopod 1 almost simple,
gnathopod 2 minutely subchelate, article 3 short.
Article 5 of pereopods 3-4 broad, slightly ex-
panded, not deeply lobate, with thin posterior
spines; dactyls of pereopods 3-7 well developed;
pereopod 5 of haustorius form, articles 2, 4, and
5 expanded, articles 4-5 with extensive facial
rows of spines; pereopods 6-7 alike, articles 5-6
weakly expanded; no pereopod with underslung
articulation.
Pleopods like urohaustonids but pleopod 3 inferior
(not 2). Epimeron 1 fully developed and setose;
epimeron 2 weakly dominant in setation and size.
Urosomites ordinary. Rami of uropod 1 styliform,
naked; of uropod 2 rod-like and spinose or with
several setae; uropod 3 of ordinary haustorioid-
phoxocephalid kind, neither ramus dominant,
both leaflike, article 2 on outer ramus very small
and poorly setose.
Telson short, broad, deeply cleft. [Sexual di-
morphism unknown.]
TYPE GENUS.?Phoxocephalopsis Schellenberg.
COMPOSITION.?Unique.
RELATIONSHIP.?Phoxocephalopsis differs from
urothoids in the antennae. It lies in position ances-
tral both to Zobrachoidae-Urohaustoriidae and
Haustoriidae, and at present appears to be the
most primitive living haustorioid known. Despite
this assertion it has several apomorphic charac-
ters, such as the simple gnathopod 1 and domi-
nant epimeron 2. One might consider Zobracho as
more primitive because of its subchelate gnatho-
pods; Phoxocephalopsis, in contrast to Zobracho, re-
tains more or less plesiomorphic uropods 1-2 and
first maxillary palp. Except for epimeron 2 and
the poorly setose maxillae and weak raker system,
it could form a plausible ancestor to the Hausto-
riidae, which have many more numerous modi-
fications in mouthparts and pereopods. Phoxoce-
phalopsis cannot be derived from any zobrachoid
or urohaustoriid without evolutionary reversal to
normalcy in uropods 1-2.
Phoxocephalopsis Schellenberg
Phoxocephalopsis Schellenberg, 1931:69 [Phoxocephalopsis zim-
meri Schellenberg, 1931, monotypy].
DIAGNOSIS.?With the characters of the family.
COMPOSITION.?The type-species and provi-
sionally P. deceptionis Stephensen, 1947.
REMARKS.?The second species of the genus, P.
deceptionis, differs from the type-species due to the
presence of 3 setae on the inner plate of maxilla
1, the weaker dominance of epimeron 2 with
epimera 1-3 all strongly setose, the parachelate
gnathopod 2 (opposed to subchelate), the poster-
oventrally produced coxa 3, and the styliform
and spinose outer ramus of uropod 2. These may
require deceptionis to be removed to a new genus.
The type-species is amplified below, by presen-
tation of new data confined mainly to informa-
tion of generic importance.
Phoxocephalopsis zimmeri Schellenberg
Phoxocephalopsis zimmeri Schellenberg, 1931:70 73, fig. 36.
MATERIAL.?Falkland Islands, Port Stanley,
No. 91, 15 Mar 1927, from mullet: female wvw,"
4.96 mm; female "y," unmeasured.
DESCRIPTION OF FEMALE (female "w," 4.96
mm).?Cephalic cheek of medium size. Eyes
small but with 5 ommatidia and pigment gran-
ules.
Length ratio of articles 1-3 of antenna 1 about
36:40:33, main armament row on article 1 con-
taining about 10 long and short setae, article 2
with both spines and setae facially, article 3 thin
and poorly setose. Article 3 of antenna 2 with 4
+ 1 short spine-setae; article 4 with dorsofacial
row of many spines in small groups, ventral mar-
gin with 3 kinds of setae (long plumose bulbar
penicillates, and short smooth spines), no glassy
bifid spines; article 5 also with dorsofacial spines.
Mandibular incisors weakly extended, right
NUMBER 360 17
and left with 4 sharp and 2-3 blunt teeth respec-
tively; right lacinia mobilis large, splayed, bifid,
apical branch sharp and smooth, proximal
branch slightly blunter and on proximal edge
pectinate almost to base, left lacinia mobilis short,
broad, pointed, with weak subapical sharp tooth
(thus technically bifid); right rakers 2, left 3, these
widely spaced, bifid and/or pectinate; molars
large, subcolumnar, triturative, choppers not
strongly distinct; palp article 1 elongate, article 2
thin, well setose on inner face, article 3 of uro-
haustoriid form but setae hooked apically, not
awned, setal formula on right = 7-1-5-1.
Mandibular lobes of lower lip intermediate in
extension between urohaustoriids and hausto-
riids, broad; inner lobes present. Inner plate of
maxilla 1 thin and leaflike, with 1 stiff but curved
medial seta, outer plate with 11 spines; palp very
short, broad, 2-articulate, article 2 slightly shorter
than 1, with 2 apical and 1 subapical setae. Plates
of maxilla 2 extending equally, subequally broad,
inner with strong oblique facial row of setae.
Maxillipeds typically urohaustoriid, inner with 3
main apical spines and inner and apical setae,
outer slightly larger than inner, medially spined;
article 2 of palp expanded, with longitudinal
facial row of setae, article 3 thinner, clavate, with
facial row of setae, dactyl small, thin, rod-shaped
or weakly clavate, apically setose. No basal balers
on mouthparts.
Coxae 1-4 progressively larger, sizes not
strongly disjunct, coxae 1-2 tapering weakly and
poorly setulose, coxa 3 blunt, scarcely produced
at posteroventral corner, there with 6-7 setae,
coxa 4 of elongate ordinary form bearing weak
posterodorsal excavation, with long posteroven-
tral and short posterior setae. Gnathopod 1 sim-
ple, gnathopod 2 subchelate but not parachelate,
otherwise of urohaustoriid form, thus bearing
long setal brushes posteriorly on articles 2-3, and
with elongate wrists, and facial apical setal row
on hands.
Pereopods 3-4 of mainly urohaustoriid form,
armament formula on pereopod 3 = 5, 3, facial
setae plus posterior spines, 6 + 5 + 1, thus differ-
ing from urohaustoriids in presence of facial setae
and long posterior spines on article 5 (instead of
spines forming pairs between face and margin);
dactyls of pereopods 3-7 well developed, simple
or bearing very weak facial tooth near apex,
dactyl of pereopod 5 lacking major spines. Per-
eopods 6-7 dominating pereopod 5, otherwise of
normal urohaustoriid form, article 2 of pereopod
5 with weak medial brush; coxa 5 lacking major
setae. Coxal gills 2-6, gill 2 dominant. Brood
plates thin.
Pleopods of generally urohaustoriid form but
pleopod 3 inferior instead of pleopod 2: coupling
hook formulas = 2 + 1,2+ 1,0+ 1, inner rami
weakly inferior on all pleopods, with basal
clothespin spine, peduncles short and broad, fa-
cial setae sparse or absent. Epimeron 2 strongly
dominant (much more so than in deceptions),
epimeron 1 well developed, both epimera 1-2
strongly setose, lacking posteroventral tooth; epi-
meron 3 smaller, poorly setose, with weak ventral
tooth near posterior margin. No dorsal wings on
pleonites 1-3.
Rami of uropod 1 styliform, sharply pointed,
inner about half as long as outer, neither with
major setae or spines apparent at low power; rami
of uropod 2 rodlike or sublinguiform, extending
equally, each with 3-4 long apical setae and 1-3
subapical setae either dorsal (on outer ramus) or
medial (on inner ramus). Armament formulas on
peduncles of uropod 1: lateral = e-E-EE-EEE,
dorsolateral = SS, medial = S-S, dorsal = E; on
uropod 2, dorsolateral = S-S-S-S, dorsal = 0,
medial = sSs. Uropod 3 ordinary, peduncle short
and broad, apicoventral margin with 6 lateral
and 5 middle spine-setae, dorsolaterally with 2
spine-setae: rami broad and fiat, article 2 on outer
ramus about 33% as long as article 1, with 2 long
and 1 short apical setae, lateral armament for-
mula = SS-SS, medial = e-ES-SS; inner ramus
as long as article 1 of outer ramus, lateral arma-
ments = S, medial = E-ES-ES-ES.
Telson broader than long, cleft almost to base,
armament formula = s-pp-s-s-p-s on each lobe,
penicillate setules listed above paired but fully
lateral.
DISTRIBUTION.?South America, south from
mouth of Rio de La Plata, and Falkland Islands,
4-40 m.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ZOBRACHOIDAE, new family
DIAGNOSIS.?Rostrum well developed (for
haustorioids), cheek poorly developed. Antenna
1 variable, article 1 short (typical) or elongate
(apomorphic), articles 2-3 progressively shorter
(typical) or elongate (apomorphic), flagella elon-
gate (typical) or not (apomorphic), articles of
peduncle weakly (typical) to strongly geniculate.
Antenna 2 of haustorius form, article 4 expanded
(pleisiomorphic) or weakly so (apomorphic), ar-
ticle 5 shorter and narrower than article 4, these
articles furnished with 1 or more longitudinal
rows of facial armaments, ventral margin of ar-
ticle 4 with at least 3 kinds of setae: (1) elongate
plumes, (2) shorter and stiffer glassy spines set in
clusters and, (3) bulbar-based penicillate setules;
flagellum longer than article 4 of peduncle. Pre-
buccal complex massive, upper lip usually domi-
nant. Mandibles bearing elongate strongly
toothed incisors, laciniae mobiles present on both
sides and unlike each other, rakers almost simple
and numerous (4 or more), molar large, strongly
extended, weakly triturative but with several
strong cusps, usually 1 of these forming accessory
chopper; palp 3-articulate, article 3 with numer-
ous outer setae, setae awned (apomorphic) or not
(typical and plesiomorphic). Lower lip with dis-
crete inner lobes, mandibular extensions of outer
lobes well developed. Maxilla 1 with uniarticulate
palp, inner plate with more than 5 setae. Maxilla
2 ordinary, inner plate with oblique facial row of
setae but poorly developed. Maxillipeds with
unexpanded bases, normally enlarged plates,
outer spinose; palp 4-articulate, article 2 ex-
panded, article 4 clavate, multisetose. No baler
lobes on maxillae or maxillipeds.
Coxa 2 small to medium, larger than coxa 1
and forming stepped intergrade between coxa 1
and coxa 3, coxa 4 dominant, coxa 3 lacking deep
posteroventral lobe. Coxal gills on segments 2-6
or 2-5. Brood plates slender.
Gnathopods feeble, subchelate, grossly alike in
proportions, wrists elongate, article 3 short. Arti-
cle 5 of pereopods 3-4 broad, slightly expanded,
not deeply lobate, with thick posterior spines;
dactyls of pereopods 3-5 well developed, those of
pereopods 6-7 variable; pereopod 5 of haustorius
form, articles 2, 4, and 5 expanded, articles 4-5
with extensive facial rows of spines; pereopods 6-
7 alike, articles 5-6 weakly expanded; no pereo-
pod with underslung articulation.
Pleopod 2 usually inferior in size, articulation,
orsetation; peduncles of pleopods not longer than
wide, inner rami inferior; coupling hooks paired
on each pleopod, usually inner rami bearing one
basal clothespin spine. Epimeron 1 moderately to
strongly developed; epimeron 2 dominant in se-
tation, often dominant in size.
Urosomites ordinary, though often furnished
with lateral teeth. Rami of uropods 1-2 lingui-
form, setose (not spinose); uropod 3 of ordinary
haustorioid-phoxocephalid kind, outer ramus
dominant, 2-articulate, peduncle short, flat, ex-
panded; rami poorly setose apically.
Telson variable in length, deeply cleft. Sexual
dimorphism weak.
TYPE GENUS.?Zobracho J.L. Barnard.
COMPOSITION.?Bumerahus, new genus; Pran-
tinus, new genus.
RELATIONSHIP.?Zobrachoids differ from uro-
thoids in the absence of a ventral cephalic cheek,
in the full development of the haustorius antenna
2, especially in the ventral armament, and in the
dominance of epimeron 2. Zobrachoids bear lin-
guiform rami of uropods 1-2, in contrast to uro-
thoids (but 1 genus of urothoid lacks rami). Pran-
tinus is furnished with a urothoid antenna 1, but
antenna 2, though not fully expanded, lacks ser-
iate ranks of spines, bears the ventral spination
diversity not typical of urothoids, and has epi-
meron 2 dominant.
Key to Genera of Zobrachoidae
1. Antenna 1 of urothoe form Prantinus, new genus
Antenna 1 of haustorius form 2
NUMBER 360 19
2. Telson elongate, rami of uropods 1-2 with many medial setae, no baso-
ventral setae Zobracho, new genus
Telson short, rami of uropods 1-2 lacking medial setae, bearing basoventral
setae Bumeralius, new genus
Zobracho J.L. Barnard
Zobracho J.L. Barnard, 1961:74.
DIAGNOSIS.?Rostrum strong. Peduncle of an-
tenna 1 short, stout, articles 2 and 3 progressively
slightly (2) and greatly (3) shortened, not genic-
ulate, both flagella elongate. Antenna 2 of haus-
torius form, article 4 expanded, article 5 small,
articles 4-5 with facial armaments, article 4 with
long ventral setae and subventral clusters of sim-
ple setae. Mandibular incisors slightly extended,
of ordinary thickness, toothed; right and left la-
ciniae mobiles not alike, right, if present, not
distinct from raker row; rakers simple, 4 or more;
molar elongate, very weakly triturative, with cus-
pidate (or spinate) accessory chopper; setae of
palp article 3 not awned, instead, minutely pec-
tinate. Mandibular lobes of lower lip well devel-
oped. Inner plate of maxilla 1 broad, multisetose,
outer plate with 11 spines; palp of ordinary
length, uniarticulate. Inner plate of maxilla 2
with weakly submarginal oblique row of setae.
Maxillae and maxillipeds lacking baler lobes.
Outer plate of maxilliped with spines; palp article
2 weakly expanded, article 3 of ordinary length,
dactyl clavate, multisetose.
Coxae 1-4 progressively larger, coxa 1 very
small but only partially hidden by following
coxae, neither coxae 3 or 4 extended posteroven-
trally; coxae 2-6 with simple gills; brood plates 4
pairs, slender.
Gnathopods small, alike, wrists elongate, hands
mitelliform, subchelate, palms well marked and
transverse. Dactyls of pereopods 3-7 distinguish-
able, those of pereopods 3-5 large, those of per-
eopods 6-7 small, dactyl of pereopod 5 bladelike,
unarmed. Article 2 of pereopods 5-7 expanded,
pereopod 5 of haustorius form, distal articles of
pereopods 6-7 not underslung, only moderately-
widened, pereopods 6-7 otherwise similar, domi-
nating pereopod 5.
Peduncles of pleopods wider than long, pleo-
pod 2 inferior, inner rami moderately shorter
than outer. Epimeron 1 strongly distinct and
multisetose, epimeron 2 dominant in size and
setation. Urosomite 1 scarcely produced and
poorly setose ventrally; peduncles of uropods 1-2
setose laterally, rami linguiform, multisetose.
Uropod 3 of ordinary form, outer ramus domi-
nant and biarticulate, article 2 with 4 or more
small or thin setae. Telson elongate, longer than
broad, deeply cleft.
DESCRIPTION.?Eyes present. Dorsolateral sur-
face of article 1 on antenna 1 furnished with
numerous elongate setae, many forming trans-
verse apical row, no longitudinal row of arma-
ments; article 2 densely setose dorsolaterally, setae
divisible into groups; article 3 poorly setose. Ar-
ticle 3 of antenna 2 short, densely setose; dorsal
margin of article 4 scarcely setose or with dense
short setae, of article 5 with dense short setae;
flagellum scarcely longer than article 4 of pedun-
cle. Calceoli present on antenna 2 in male.
Right lacinia mobilis indistinguishable from
raker row, left slender, bifid. Lower lip lacking
cones. Inner plate of maxilla 1 with long setae on
medial, apical and lateral margins; several spines
on outer plate weakly bifid. Inner plate of max-
illiped with 3 stout apical spines (type-species
unknown).
Coxae 1-2 both poorly setose (type-species) or
coxa 2 strongly setose. Gill 2 dominant, gill 5
much larger than posterior lobe of coxa 5, gill 6
very small. Gnathopods 1-2 lacking surficial but-
ton combs (see Urohaustonus).
Pereopods 3-4 lacking secondary facial row of
spines on article 5, except most distal spine on
margin with facial partner.
Uropods 1-2 lacking huge dorsal setae on pe-
duncles; outer rami setose both laterally and me-
dially, no basoventral setae.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TYPE-SPECIES.?Zobracho canguro J.L. Barnard,
1961 (original designation).
COMPOSITION.?Unique.
RELATIONSHIP.?Of the genera bearing haus-
torius-like pereopod 5 combined with uniarticu-
late first maxillary palp, this genus appears to
stand in the primitive position. This is also re-
flected in the large rostrum and broad, unmodi-
fied, multisetose inner plate on maxilla 1. Genera
such as Phoxocephalopsis and those in the Urothoe
group retain the plesiomorphic maxilla 1 with 2-
articulate palp and most of these also have at
least uropod 1 with nonlinguiform rami. Zobracho
would appear to be a very close antecedent to
Bumeralius.
Zobracho canguro J.L. Barnard
Figures 2-6
Zobracho canguro J.L. Barnard, 1961:75, figure 43.
DESCRIPTION OF MALE (male "m," 6.25 mm).?
Head about as long as wide, rostrum about 44%
as long as remainder of head. Primary flagellum
of antenna 1 with 11 articles, articles 3-10 with
long aesthetascs in groups of 3 usually; accessory
flagellum 10-articulate. Article 3 of peduncle on
antenna 2 with about 10 facial setae; facial for-
mula of thin spines on article 4 = 6-9-12-many;
on article 5 = many; distomedial face of article 5
with 2 (1) + ?1 rudimentary large flask-shaped
calceoli, each of many compartments; flagellum
with 1 I articles, first 3 bearing similar calceoli.
Right and left mandibular incisors with 3 teeth,
right rakers 8 (including lacinia mobilis) plus 2
rudimentaries, left 7 plus 2 rudimentaries, chop-
per on molar with 3 cusp-spines; article 1 of palp
scarcely elongate, article 3 slightly shorter than
article 2, latter of medium setosity, spine formula
on right and left article 3 = 9-1-6-1. Inner plate
of maxilla 1 with 2 short appressed apicomedial
setae (marking apex of longitudinal facial ridge),
7 long laterals and 3 long medials; outer plate
with 1 1 spines; palp with 2 stout and 1 thin apical
setae. Inner plate of maxilliped with 11 submar-
ginal medial setae continued in crescent to api-
colateral margin, 3 stout apical teeth, most distal
disjunct, medial margin of outer plate with short
thin spines in tandem, then notch bearing seta, 1
more spine on protrusion, then several thin setae
in tandem more distally, apex with 5 medium
setae; article 2 of palp with apical group of thin
facial setae, article 3 with setal ranks facially,
lateral margin with group of 4 setae.
Coxa 1 very short, wide, hatchet-shaped, pos-
teroventrally extended, with 1 long posteroven-
tral seta and many marginal setules; coxa 2 much
larger than coxa 1, elongate, weakly truncate
apically, bearing 9 long apical setae; coxa 3
slightly larger than coxa 2, with 10 long apical
setae (some medium in length); coxa 4 with about
8 medium ventral setae, posterior margin with
about 19 medium and short mixed setae; coxa 5
distinctly bilobed. Gnathopods 1 and 2 similar,
feeble, subchelate; hands with short, almost trans-
verse palms, slightly humped on gnathopod 2;
article 3 very short, with numerous posterior se-
tae; wrists much longer than hands (relatively
longer on gnathopod 2 than gnathopod 1), both
strongly setose posteriorly and with medial facial
setae in groups; article 6 with well-defined lateral
facial crescent of long strong setae distad.
Setal and spine formulas on pereopod 3 = 8,
12, 2-1-1-1, 6 + 4 + 1; on pereopod 4 = 6 (plus
4 on anterior margin), 11, 2-1-1-1, 6 + 5 + 1,
article 4 especially small; posterior margin of
article 5 on pereopods 3-4 with groups of serra-
tions. Article 2 of pereopod 5 densely setose pos-
teriorly, of pereopod 6 of lesser density, of pereo-
pod 7 scarcely setose; pereopod 5 with brush of
mediofacial setae posteroventrally, dactyl long,
naked (except for regular basofacial setule),
bladelike, with small anterodistal notch and
tooth, dactyls of pereopods 6-7 shorter than ad-
jacent spines; dactyls of pereopods 3-4 also with
tooth and notch. Medial apex of article 6 on
pereopod 7 softly castellate.
Pleopod 2 inferior; peduncles of all pleopods
with sets of 2-5 setae anteriorly, spine formulas
all = 2 and 1, coupling hooks like those of Uro-
haustonus; each inner ramus with 2 basal clothes-
pin spines; segmental formulas = 22-15, 20-15,
NUMBER 360 21
FIGURE 2.?Zobracho canguro J.L. Barnard, male "m," 6.25 mm (h = female "h," 6.26 mm; k
juvenile "k," 4.36 mm).
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
D5
FIGURE 3.?Zobracho canguro J.L. Barnard, male "m," 6.25 mm (h = female "h," 6.26 mm; Y ?
penial process; Z= brood lamella).
NUMBER 360 2:3
FIGURE 4.?Zobracho canguro J.L. Barnard, male "m," 6.25 mm (h = female "h," 6.26 mm; Z
brood lamella).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 5.?Zobmcho canguro J.L. Barnard, male "m," 6.25 mm (h ? female "h," 6.26 mm: Z =
gill)-
NUMBER 360 25
FIGURE 6.?Zobracho canguro J.L. Barnard (j = juvenile " j , "
4.2 mm; r = male "r," 7.0+ mm; w = female "w," 8.0+
mm).
20-16; basal setal formulas = 12-2-3-5, 7-2-2-
4, 9-2-2-5.
Epimeron 1 fully developed, extended in adze-
form posteroventrally, bearing posterior row of 5
setae closely crowded, 1 seta near posteroventral
corner and 2 other facial setae in horizontal row
near posterior corner; epimeron 2 sharply pro-
duced posteroventrally, ventral margin concave,
posterior margin and corner with 5 large setae
and 4-5 setules, ventral margin with 14 thin
setae; epimeron 3 with sharp posteroventral
tooth, bulging posteriorly above sinus, margin
setulate.
Lateral margin of peduncle on uropod 1 with
8-9 setae and 2 apical setules, of uropod 2 with
6 setae and 2 setules, no dorsal setae; peduncles
with 4 and 3 groups of 1-4 setae each on medial
margin; peduncles of uropods 1-3 densely packed
with glandular tissue; outer rami of uropods 1-2
each with 3-4 lateral setae (paired on uropod 2),
apices with 4, medial margins with 7 setae; inner
rami each with 1 lateral seta, apices with 4,
medial margins with 8 and 7 groups of 1-3 setae
each; these setae much weaker than in Urohaus-
tonus; inner rami slightly shorter than outer rami.
Uropod 3 weakly elongate, peduncle naked lat-
erally, with 2 apicolateral setules, with pair of
small setae medially, 1 apicomedial spine, and
row of 5 small setae on ventral margin; inner
ramus reaching M. 80 on article 1 of outer ramus,
with setae medially, apically, and apicolaterally;
article 1 of outer ramus with pairs of lateral setae
but simply setose medially, article 2 elongate,
0.42, bearing 5 weak apical setae (some long,
others short). Urosomal glands other than those
in peduncles of uropods not apparent.
Telson elongate, deeply cleft, slightly broad-
ened at base, setal formula variable, = 2P, 0-1,
P, 2-3, apical setae variable in placement.
Cuticle with very sparse bulbar setules and
extremely faint but dense pebbling.
DESCRIPTION OF FEMALE (female "h," 6.26
mm).?Like male except as follows: Eyes smaller
than in male. Primary flagellum of antenna 2
with 9 articles, accessory flagellum with 8, aesth-
etascs shorter than in male and only 1 per article.
Several facial setae on article 4 of antenna 2
thicker than in male; flagellum of antenna 2 with
11 articles. Right mandibular rakers 8 plus 3
rudimentaries, left 7 plus 3 rudimentaries, for-
mula on palp article 3 = 8-1-6-1. Inner plate of
maxilla 1 with 2-3 medial setae, 2 apical, 5-6
lateral setae. Inner plate of maxilla 2 with 11
setae in oblique row (including 1 medial seta).
Coxae 1-2 like male; coxae 3-4 slightly more
elongate (illus.). Hand of gnathopod 2 stouter
and less tapering than in male (illus.).
Formulas of pereopod 3 = 6, 10, 2-1-1-1, 6 +
5 4- 1; of pereopod 4 = 5, 10, same, same. Setae
on coxae and article 2 of pereopods 5-6 longer
than in male.
Epimeron 2 with 12 ventral setae, 3 at poster-
oventral corner (as in male) and 3 above that
group with dorsalmost very discontiguous.
Uropods 1-2 similar to male, generally with 1
fewer seta or group of setae in each category, with
only 1 apicolateral setule on peduncles. Peduncle
of uropod 3 with 3-4 apicolateral setules, ventro-
medial spine and setae longer than in male, apical
setae on article 2 of outer ramus longer than in
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
male, rami otherwise slightly less setose than in
male.
DESCRIPTION OF JUVENILE (juvenile tkk," 4.36
mm).?Eyes very weak, with only 5-6 ommatidia
each. Right mandibular rakers 7 plus 2 rudimen-
taries, left with 6 plus 2 rudimentaries; molars
with only 2 distinct cusps on choppers; each palp
article 3 = 5-1-5-1. Epimeron 1 with 1 posterior
medium seta and 1 setule at corner. Epimeron 2
with 3 ventral setae, corner with 1 large seta,
indented, and 1 setule, posterior margin with 1
large, 1 small seta. Telson with either 1 or 2 long
setae on each side.
OBSERVATIONS.?All lateral setae of peduncle
but few medial setae on rami of uropod 2 plu-
mose.
SPECIMENS FROM NEW SOUTH WALES.?Much
larger than those from Victoria, largest in the
collection measuring nearly 10 mm in length.
In haustorioids generally, and particularly in
zobrachoids, a small increase in length, as meas-
ured from tip of rostrum to base of telson, results
in a disproportionately large increase in the mas-
siveness of the individual; and in a specimen 9
mm long the appendages are so very much larger
than those of the biggest in the Victorian collec-
tion (6.93 mm) that it appears, at first sight,
unlikely to belong to the same species. Careful
examination of a number of dissected specimens,
however, has revealed no differences that could
not be ascribed to the greater size of the New
South Wales specimens. Setae and spine-bearing
appendages in general are, as might be expected,
more setose or spinose; but a significant exception
to this is coxa 1 which, in juveniles and adults
alike, bears, as in Victorian specimens, a single
apical seta. Subtleties of coxal shape, emphasized
and accentuated by larger size, contribute to the
appearance of dissimilarity; but small differences
between male and female article 2 of pereopods
5-7 are the same as in Victorian specimens.
The most striking difference between New
South Wales and Victorian specimens is found in
the "'cusp" of epimeron 3; long, stout, and straight
in the New South Wales collection, but short and
up-turned in every Victorian specimen except
female "p," where it has become straight.
Examination of a single sample (through the
courtesy of Dr. Deborah Dexter, 1981) from New
South Wales, containing individuals at all stages
of development, suggests the possibility that the
Victorian specimens may not have reached the
terminal stage.
The epimeral cusp in New South Wales juve-
niles is small and inturned, as in Victorian spec-
imens; article 2 of the outer ramus of uropod 3 is
much longer, relative to article 1, than it is in the
adult, and the inner ramus is considerably shorter
than article 1 (a typical haustorioid juvenile char-
acter). The number of coxal setae in juveniles is
small and increased with increasing size. But
juveniles of about 5 mm have 4-5 setae on coxa
2, though only 1 on coxa I. In the largest female
in the sample, 9 mm, with setose brood plates
(assumed terminal), article 2 of the outer ramus
of uropod 3 is little more than 0.25 as long as
article 1, and the inner ramus reaches beyond the
apex of that article.
The condition of uropod 3 (illus.) and the small
upturned epimeral cusp of the holotype suggests
that it is not a terminal male. Female "p," with
setose brood plates has, compared with the holo-
type, a shorter (0.33) article 2 of the outer ramus
of uropod 3 and a longer inner ramus; the cusp
is straight though short. It is closer in general
form to the large New South Wales specimens,
but, judging by uropod 3, it is possible that this,
too, may be subterminal.
ILLUSTRATIONS.?Left mandibular palp article
2 not flattened. Lateral face of hand on gnatho-
pod 1 like gnathopod 2. Fully delineated pereo-
pod 5 of Figure 4 enlarged; other peropod 5,
lacking setae, magnified to same degree as per-
eopods 6-7 and marked with positions of medial
setae.
HOLOTYPE.?See J.L. Barnard (1961); coxa 1
with 2 apical setae and coxa 2 with only 2 apical
setae; the small count on coxa 2 is bridged by
various intermediate specimens.
VOUCHER MATERIAL.?CPBS 35S/1: male
"m," 6.25 mm (illus.); WPBES 1726/3: female
NUMBER 360 27
"h," 6.26 mm (illus.); CPBS A4/5: female "p,"
6.93 mm; juvenile "k," 4.36 mm (illus.); AM SBS
Transect 2.09.03.05 (2 specimens); AM P.24856:
female "w," 8.0+ mm (illus.); AM P.24857: male
"r," 7.01+ mm (illus.); AM P.23375: juvenile " j , "
4.2 mm (illus.).
REMARKS.?Specimens from New South Wales
may merit specific status. Additional collections
from Victoria containing adults could clarify the
relationship.
MATERIAL.?CPBS, 11 samples from 9 stations
(16); WPBES, 1 sample (1); AM HDWBS and
SBS, 43 samples from 18 stations (158).
DISTRIBUTION.?Great Australian Bight, 875 m;
Victoria, Western Port, confined to channel sta-
tions (not taken in Port Phillip); NSW, 16 km
south to 3 km north of Hunter River, intertidal
to 28 m, sandy beaches and off shore. Fine to
coarse sand.
Bumeralius, new genus
DIAGNOSIS.?Rostrum weak. Peduncle of an-
tenna 1 short, stout, articles 2 and 3 of peduncle
progressively shortened, geniculate between arti-
cles 1 and 2, both flagella moderately long. An-
tenna 2 of haustorius form, article 4 expanded,
article 5 small, articles 4-5 with facial armaments,
article 4 with long ventral setae and subventral
clusters of bifid setae. Mandibular incisors not
extended, thick, toothed; right and left laciniae
mobiles not alike; rakers simple, 3 or more; molar
elongate, weakly triturative, with cuspidate ac-
cessory chopper; setae of palp article 3 awned.
Mandibular lobes of lower lip well developed.
Inner plate of maxilla 1 slender, multisetose, outer
plate with 11 spines, palp short, uniarticulate.
Inner plate of maxilla 2 with fully submarginal
oblique row of setae. Maxillae and maxillipeds
lacking baler lobes. Outer plate of maxilliped
with spines, palp article 2 expanded, article 3
elongate, dactyl clavate, multisetose.
Coxae 1-4 progressively larger, coxae 1-2 small
but only partially hidden by following coxae,
neither coxae 3 or 4 extended posteroventrally;
only coxae 2-5 with simple gills; brood plates 4
pairs, slender.
Gnathopods small, almost alike, wrists elon-
gate, hands mitelliform, subchelate, palms well
marked. Dactyls of pereopods 3-7 distinguish-
able, those of pereopods 3-5 large, those of per-
eopods 6-7 small, dactyl of pereopod 5 bladelike,
unarmed. Article 2 of pereopods 5-7 expanded,
pereopod 5 of haustorius form, distal articles of
pereopods 6-7 not underslung, only moderately
widened, pereopods 6-7 otherwise similar, domi-
nated by pereopod 5.
Peduncles of pleopods wider than long, pleo-
pod 2 inferior, inner rami moderately shorter
than outer. Epimeron 1 scarcely differentiated,
epimeron 2 dominantly setose. Urosomite 1
scarcely produced and poorly setose ventrally.
Peduncles of uropods 1-2 setose laterally, rami
linguiform, bearing only 1 apical seta each except
for outer ramus of uropod 2. Uropod 3 of ordinary
form, outer ramus dominant and biarticulate,
article 2 with 2 apical setae. Telson short, broader
than long, deeply cleft.
DESCRIPTION.?Eyes indistinct or absent, ocular
ganglia visible. Dorsolateral surface of article 1
on antenna 1 furnished with numerous elongate
setae, not organized into distinct longitudinal
row; article 2 densely setose dorsolaterally, groups
indistinguishable; article 3 poorly setose. Article
3 of antenna 2 short, densely setose; dorsal margin
of article 4 scarcely setose, of article 5 strongly
setose; flagellum longer than article 4 of peduncle.
No calceoli.
Right lacinia mobilis thin, narrow; left
broader, both bifid. Lower lip with 1 cone on
each lobe. Inner plate of maxilla 1 with mostly
short apical setae, excavate medial margin
bounded proximally by large seta; most spines on
outer plate weakly bifid and/or marginally pec-
tinate. Inner plate of maxilliped with 3 stout
apical spines.
Coxae 1-2 strongly setose. Gill 2 dominant, gill
5 much larger than posterior lobe of coxa 5.
Gnathopod 1 bearing surficial button-combs (see
"Urohaustonus," Figure 21 QGlt).
Pereopods 3-4 with extensive secondary facial
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
row of spines on article 5 in addition to normal
posterior spines.
Uropods 1-2 lacking huge dorsal setae on pe-
duncles; outer rami of both pairs usually with
group of basomedial setae, medial margin of
peduncle on uropod 2 almost naked.
TYPE-SPECIES.?Bumeralius buchalius, new spe-
cies.
COMPOSITION.?Unique.
RELATIONSHIP.?Bumeralius is considered to be
descended from a morphotype like Zobracho J.L.
Barnard (1961), as shown by the shortened telson
and sparser and more specialized setation on the
rami uropods 1-2. Of lesser importance is the
narrowed inner plate of maxilla 1 with specialized
setation pattern, the shorter palp of maxilla 1, a
slightly stronger distinction between the hands of
gnathopods 1 and 2, the lesser dominance by the
epistomal part of the prebuccal complex, and the
smaller rostrum.
Bumeralius buchalius, new species
FIGURES 7-12
DESCRIPTION OF HOLOTYPE MALE (male "o,"
5.60 mm).?Head about 85% as long as wide,
rostrum about 21% as long as remainder of head.
Primary flagellum of antenna 1 with 8 articles,
first 5 each with long aesthetasc, accessory flagel-
lum 5-articulate. Article 3 of peduncle on antenna
2 with about 14 facial setae; facial formula of
thin setae on article 4 = 9-10-many, on article 5
= many; flagellum with 10 articles, at least 2
setae on each article elongate; no calceoli ob-
served on articles of peduncle or on flagellum.
Right incisor with at least 3 teeth, left with 2,
right rakers 3, left 5 plus I rudimentary, chopper
on molar with 2-3 cusps; article 1 of palp weakly
elongate, articles 2 and 3 subequal in length,
article 2 sparsely setose, spine formula on right
article 3 = 16-1-9-1, on left = 15-1-8-1. Inner
plate of maxilla 1 with 2 short apical setae, 2
longer laterals, 1 longer apicomedial, 1 elongate
mediobasal; outer plate with 11 spines; palp with
2 apical setae and 1 disjunct apicomedial seta.
Inner plate of maxilliped with 6 submarginal
medial setae, medial margin of outer plate with
ragged mixture of spines and groups of small
setae, apex naked, apicolateral margin with 1
seta; article 2 of palp with row of 8 facial setae,
article 3 with row of 5 (or 6 counting 1 near apex)
facial setae.
Coxa 1 short, wide, blunt, bearing 23 small
setae densely crowded; coxa 2 of skewed pyriform
shape, with distoventral extension, anterior mar-
gin with 20 short to medium thin setae, ventral
apex with 8 longer plumes, posterior margin with
3 shorter plumes; coxa 3 with 6 ventral and 11
posterior plumes; coxa 4 with 7 ventral and about
32 posterior setae of mixed sizes and plumosities;
coxa 5 very short, unlobed or scarcely lobed.
Gnathopods with posterior setal brushes on arti-
cles 2-3; palm of gnathopod 1 slightly oblique,
that of gnathopod 2 transverse, margins distinc-
tive (see illustrations); hand of gnathopod 2
smaller than that of gnathopod 1.
Setal and spine formulas on pereopod 3 = 8,
13, 8 facial + 5 posterior, 7 + 4 + 1; on pereopod
4 = 8, 10, 7 facial 4- 6 posterior, 7 + 4 4- 1;
margins of articles smooth; dactyls of pereopods
3-4 lacking subapical ornament. Article 2 of per-
eopods 5-7 densely setose posteriorly, that of
pereopod 5 with posteroventral brush of mediofa-
cial setae, of pereopod 7 with posterodorsal hook;
dactyl of pereopod 5 long, almost naked, blade-
like, with small anterodistal notch or tooth, dac-
tyls of pereopods 6-7 not longer than adjacent
spines; medial apex of article 6 on pereopod 7
with pectinate invagination.
Pleopod 2 inferior; peduncles of dominant
pleopods with rows of 4-5 and upwards to 10
long setae, spine formulas of pleopods 1-3 = 2
and 2, 2 and 0, 2 and 2, coupling hooks like those
of Urohaustorius; each inner ramus with basal
clothespin spine; segmental formulas of pleopods
= 25-16, 18-12, 22-19 (inner proliferate); basal
setal formulas = 21-2-3-3, 13-0-1-1, 13-(3-4)-
4-5.
Epimeron 1 obsolescent, bearing 1 posteroven-
tral setule; epimeron 2 produced posteroventrally
like adze, with subventral horizontal row of 8-9
NUMBER 360 29
.
FIGURE 7.?Bumeralius buchalius, new species, holotype, male "o," 5.60 mm (p = female "p,
4.43 mm; Y? left lacinia mobilis).
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 8.?Bumeralius buchalius, new species, holotype, male "o," 5.60 mm (p = female "p,
4.43 mm; Z= gill).
NUMBER 360 31
E
FIGURE 9.?Bumerahus buchalius, new species, holotype, male "o," 5.60 mm.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ZP3
FIGURE 10.?Bumeralius buchalius, new species, holotype, male "o," 5.60 mm ( Y = pleopod; Z =
gill)-
NUMBER 360 33
M
KP5
FIGURE 11.?Bumeralius buchalius, new species, unlabeled = juvenile " j , " 2.97 mm (u = juvenile
"u," 2.54 mm).
setae, posterior submarginal row of 4-5 setae,
posterior margin with setule; epimeron 3 with
posterior hump strongly ventrad, posterior and
ventral margins weakly sinuous, posteroventral
corner softly rounded, margins bearing only few
setules.
Lateral margin of peduncle on uropod 1 with
9 setae, extending to venter, of uropod 2 with 4,
not extending to venter and with naked gap to
apex bearing 2 setules, no dorsal setae; medial
margin of uropod 1 densely setose, of uropod 2
with 1 seta; outer rami of uropods 1-2 each with
elongate apical plume, inner rami with shorter
stiffer seta, outer ramus of uropod 2 with 2 lateral
plumes, all other ramal margins naked; outer
ramus of uropod 1 with 8 long plumes in subbasal,
ventral, and medial cluster, uropod 2 with 2 in
same position; inner rami about 70% as long as
outer. Uropod 3 short, peduncle with 3 large
lateral setae, 1 medial setule, dense apicoventral
setae; inner ramus reaching article 1 apex of outer
ramus, with 7 medial, 3 apical, and 1 lateral
setae; article 1 of outer ramus with 6 lateral setae
in 5 groups, mostly crowded distally, medial mar-
gin with 3 widely spread setae, article 2 elongate,
0.48, bearing 2 long apical setae and 1 apicome-
dial setule. Urosomal glands not apparent.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 12.?Bumeralius buchalius, new species, juvenile " j , " 2.97 mm.
Telson very broad, deeply cleft, ungaping, with
moderately developed lateral alae, setal formula
= 1-2, 2P, 3-4, P, 2-3.
DESCRIPTION OF FEMALE.?Female "p," 4.43
mm: Antenna 1 like male but long setae fewer,
primary flagellum with 8 articles, accessory with
7. Antenna 2 like male but facial setae slightly
fewer, flagellum with 9 articles. Right mandible
with fourth raker rudimentary, left with fifth
raker rudimentary; left lacinia mobilis with 4
teeth (illus.); spine formula on right palp article
3 = 12-1-7-1, on left = 13-1-6-1. Setae on inner
plate of maxilla 1 more elongate than in male
(illus.). Outer plate of maxillipedal palp with
only 4 medial spines plus setal groups, article 2 of
palp with 6 facial setae, article 3 with either 5 or
NUMBER WO 35
6 facial setae. Setae of coxa 3 more numerous
than in male. Brood plates 4 pairs, narrow and
setose. Setae ofgnathopod 1 fewer than in male.
Setal and spine formulas on pereopod 3 = 6, 8, 5
facial + 5 posterior, 5 + 3 + 2!; on pereopod 4
= 5, 6-7, 5 facial + 6 posterior, 5 + 3 + 1. Article
2 of pereopod 5 with only 2 anterodorsal setae,
article 4 with only 2 spines in row, other spines
almost as numerous as in male; rows of spines on
article 5 of pereopod 6 shorter and sparser than
in male, dactyl thinner and longer.
Pleopods differing from male as follows: seg-
mental formulas = 20-12, 16-10, 20-16, basal
setal formulas = 15-2-3-4, 11-0-1-1, 12-2-3-5.
Epimeron 2 with 7 ventral, 3 upper oblique long
setae; epimeron 3 with 2 posteroventral and 1
anteroventral setules.
Uropod 1 with 8 lateral peduncular setae, outer
ramus with 6 basomedial setae; uropod 2 with 3
lateral peduncular setae and 2 apical setules, 1-2
medials, outer ramus with 1-2 lateral setae, 1
basomedial; peduncle of uropod 3 with 2 lateral
setae, otherwise see illustration. Telsonic setal
formula = 2P, 3-4, P, 3 (see illustration).
Female "a," 4.89 mm: Outer ramus of left
uropod 3 with 3 apical setae on article 2, third
seta actually apicolateral. Telsonic setal formula
= 2P, 3, P, 2.
ILLUSTRATIONS.?One spine on face of article 5
on pereopod 3 unemergent, marked in dots; all
setae of uropod 1 plumose but apicalmost setae
of rami only plumose terminally; some setae of
uropod 2 poorly plumose.
DESCRIPTION OF JUVENILE.?Juvenile " j , " 2.97
mm: Like adult, but with the juvenile charac-
teristics of much fewer spines and setae. Head
about 87% as long as wide, rostrum about 24% as
long as remainder of head. Primary flagellum of
antenna 1 with 5 articles, accessory flagellum
with 4-5 articles. Facial formula of thin setae on
article 4 of antenna 2 = 3-15, on article 5 = 8;
flagellum with 7 articles.
Right incisor weakly bifid, left with 3 teeth;
right rakers 3, left 4; chopper on molar with 2
cusps and hump; left lacinia mobilis with 3 teeth;
article 2 of palp with 1 seta, spine formula on left
palp article 3 = 6-1-4-1. Cones of lower lip very-
large. Inner plate of maxilla 1 with 2 short apical
setae, no laterals, 1 longer apicomedial, 1 elongate
mediobasal; outer plate with 9 spines. Inner plate
of maxilla 2 with 5 setae in oblique row. Inner
plate of maxilliped with 3 apical stout spines and
4 submarginal medial setae, outer plate with 3
medial spines and groups of 2, 3, and 2 setae,
apex naked; article 2 of palp with row of 4 facial
setae, article 3 with row of 3 facial setae, article
4 with 2 apical setae, 1 dorsal, and 3 small setae
on inner margin.
Coxa 1 with 6 setae, coxa 2 short, deeply incised
posteriorly, bearing 5 anterior setae, 3 apical (long
and short) and 1 posterior marginal; coxa 3 with
9 setae, coxa 4 with 12 setae. Gills like adult.
Gnathopods approximately like adult but less
setose, article 5 slightly thinner, medial face of
hand with only 1 enlarged seta near dactyl, palm
of gnathopod 1 with only 3 small spinules, of
gnathopod 2 with only 5 small spinules.
Setal and spine formulas on pereopod 3 = 2, 2,
1 facial + 3 posterior, 3 + 1 + 1; pereopod 4 =
same. Pereopod 5 lacking brush of mediofacial
setae on article 2, dactyl lacking any setae, per-
eopods 5-7 otherwise similar to adult, dactyl of
pereopods 3-4 with weak subapical acclivity-
tooth; medial apex of article 6 on pereopod 7
only with pectinate invagination; article 2 with
posterodorsal hook.
Peduncular spine formulas of pleopods 1-3 =
2 and 0, 2 and 0, 2 and I, each inner ramus with
basal clothespin spine; segmental formulas of
pleopods 1-3 = 12-8, 10-5, 13-8; basal setal
formulas =10-1-1-2, 5-0-1-1, 7-1-1-2, pleopod
2 scarcely inferior.
Epimeron 1 obsolescent, bearing 1 posteroven-
tral setule; epimeron 2 produced posteroventrally
like adze, with ventral row of 2 setae, posterior
row of 2 setae, posterior margin with 1 setule;
epimeron 3 with hump and posteroventral corner
amalgamated as broadly rounded outline.
Lateral margin of peduncle on uropod 1 with
6 setae extending to venter, on uropod 2 with 3
not extending to venter and with gap to naked
apex, medioventral face of peduncle with 6 setae.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
or uropod 2 with 1 seta; outer rami of uropods 1-
2 each with elongate apical plume, inner rami
with shorter stiffer almost naked seta, all rami
otherwise naked marginally; outer ramus of uro-
pod 1 with 2 long basal plumes, of uropod 2 with
1 in same position, inner rami about 20% as long
as outer. Uropod 3 somewhat elongate, peduncle
with 1 large lateral seta, 1 setule medially, sparse
(5-6) apicoventral setae, inner ramus with 3 setae,
article 1 of outer with 1 apicolateral and 1 mid-
medial setae, article 2 elongate, bearing 2 long
apical setae and 1 apicomedial setule. Urosomal
glands not apparent.
Telson very broad, deeply cleft, ungaping, with
moderately developed lateral alae, setal formula
= 2P, P, 0-1.
Juvenile "u," 2.54 mm: Epimeron 2 with only
1 posterior seta and setule, 1 facial seta on ventral
row; epimeron 3 with only 1 setule (posteroven-
tral corner); peduncle of uropod 1 with 4 lateral
setae, outer rami of uropods 1-2 each with 1
basomedial seta, peduncle of uropod 2 with 1
lateral seta; peduncle of uropod 3 with only 3
ventrodistal setae, article 1 of outer ramus with
only 1 seta situated apicolaterally, inner rami
with 4 setae (2 apical, 2 basomedial); telson illus-
trated.
Juvenile "h" (larger size): Inner ramus of
uropod 1 about 34% as long as outer, of uropod
2 about 55% as long as outer ramus.
HOLOTYPE.?NMV, male "o," 5.60 mm (illus.).
TYPE-LOCALITY.?WPBES 1715, Western Port,
Victoria, 10 Jan 1974, intertidal, coarse sand.
VOUCHER MATERIAL.?AM MER 3D, Merim-
bula, NSW: female "p," 4.43 mm (illus.); juvenile
" j , " 2.97 mm (illus.), juvenile "u," 2.54 mm (il-
lus.); MER 37B: juvenile "h," larger size; CPBS
C6/1: female "a," 4.89 mm, and young female
tvg," 3.70 mm, with half-grown, unsetose brood
plates.
REMARKS.?The condition of rami on uropods
1-2 in large juvenile "h" seems to confirm the
transition from small juveniles to adults. Gener-
ally the inner rami of uropods 1-2 in small juve-
niles is about 20%- of the length of outer rami and
in adults about 70%. The rami of juvenile "h,"
34%) and 55% respectively on uropods 1-2 seems
to bridge this gap.
MATERIAL.?CPBS, 1 sample (1); WPBES, 1
sample (1); AM Merimbula, (Day samples) 2
samples (9).
DISTRIBUTION.?Victoria and New South
Wales, intertidal.
Prantinus, new genus
DIAGNOSIS.?Rostrum short and broad but
head extended strongly anteriad from antennal
notch. Peduncle of antenna 1 elongate, thin, ar-
ticles 2 and 3 about 70%. and 45%; as long as
article 1, strongly geniculate, both flagella short,
scarcely longer than article 3 of peduncle. An-
tenna 2 of weak haustorius form, article 4 weakly
expanded but bearing 3 kinds of armaments, long
ventral setae, subventral clusters of non-bifid but
apically sheathed spines and facial armament
row; article 5 much smaller than article 4. Aesth-
etascs simple. Mandibular incisors extended,
toothed; right and left laciniae mobiles not alike;
rakers serrate, numerous on each mandible; molar
small, thin, extended, with 2 main and 2 subsid-
iary apical cusps plus whiplike seta; setae of palp
article 3 not awned, apically hooked. Mandibular
lobes of lower lip well developed. Inner plate of
maxilla 1 of medium size, moderately setose,
outer plate with 10 spines, palp short, uniarticu-
late. Inner plate of maxilla 2 with weakly sub-
marginal row of setae. Maxillae and maxillipeds
lacking baler lobes. Inner plate of maxilliped
ordinary in female, unusually small in male; outer
plate spinose; palp article 2 weakly expanded,
article 3 not extraordinarily elongate, slightly
expanded apically, dactyl unguiform, elongate,
bearing apical nail and subsidiary setae.
Coxae 1-4 progressively larger, coxae 1-2
small, subequal in size, coxa 3 shaped like tear
drop, not greatly enlarged, and not covering an-
terior coxae, coxa 4 not greatly enlarged, subov-
ate; coxae 2-5 with simple gills; brood plates 4
pairs, slender.
Gnathopods small, grossly alike, wrists elon-
gate, hand somewhat smaller, more mitelliform,
NUMBER 360
subchelate, but palm more transverse on gnath-
opod 2 than on gnathopod 1. Dactyls of pereo-
pods 3-7 distinguishable, those of pereopods 3-5
large, those of pereopods 6-7 very small; dactyl
of pereopod 5 lacking spines. Article 2 of pereo-
pods 5-7 expanded less strongly on pereopods 6-
7 than on pereopod 5; latter of haustorius form;
distal articles of pereopods 6-7 not underslung,
only moderately widened; pereopods 6-7 other-
wise similar, not dominating pereopod 5.
Peduncles of pleopods wider than long, pleo-
pod 2 slightly inferior, inner rami shorter than
outer. Epimeron 1 poorly differentiated, epime-
ron 2 dominantly setose. Urosomite 1 weakly
produced and strongly setose ventrally. Rami of
uropods 1-2 styliform, each bearing only 1 apical
plumose seta; peduncle of uropod 1 in male
strongly setose laterally, of uropods 1-2 in female
weakly spinose apicolaterally. Uropod 3 of ordi-
nary form; outer ramus biarticulate and domi-
nant in female, inner ramus dominant in male;
article 2 of outer ramus with 2 large and 1 minute
apical setae. Telson short, broader than long,
deeply cleft.
DESCRIPTION.?Eyes weak to indistinct in fe-
male and juveniles, well developed in male, ocu-
lar ganglia visible. Dorsolateral surface of article
1 on antenna 1 furnished with small poorly orga-
nized group of setae; article 2 moderately setose
dorsolaterally; article 3 poorly setose. Article 3 of
antenna 2 short, moderately setose, flagellum
much longer than article 4 of peduncle. No cal-
ceoli yet observed.
Right lacinia mobilis bifid, left broader and
trifid. Lower lip with inner lobes almost com-
pletely fused together, cones on outer not appar-
ent. Inner plate of maxilla 1 with apical setae
both medial and lateral; several spines on outer
plate either bifid or with accessory spine. Inner
plate of maxilliped with 3 stout apical spines.
Coxa 1 poorly setose, coxae 2-4 moderately
setose. Gill 4 dominant, gill 5 larger than posterior
lobe of coxa 5. Gnathopod 2 lacking surficial
buttons (see "Urohaustorius").
Pereopods 3-4 lacking extensive secondary fa-
cial rows of spines on article 5.
Uropods 1-2 in male with sparse dorsal setae,
medial margins of peduncles strongly setose in
both sexes.
TYPE-SPECIES.?Prantinus talanggi, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?Prantinus forms an evolution-
ary cluster of its own because of the combination
of a urothoe antenna 1 with a weakly expanded
antenna 2 of the haustorius form. It has many
similarities to Zobracho and could be interpreted
as being a descendent in which antenna 1 has
diverged and the telson become short; otherwise
it resembles Zobracho in the head and subchelate
gnathopods. The rami of uropods 1-2 are unique
to the greater haustorioid complex of genera,
though an intermediate stage between the lingui-
form kind of rami in Urohaustorius and those of
Prantinus are present in Bumerahus. The latter
genus also could be visualized as an intergrade
between Zobracho and Prantinus because of similar-
ities to Prantinus in apical armament on article 2
of the outer ramus on uropod 3, the setose con-
dition of the inner plate on maxilla 1, the shapes
of coxae, mandibular molar, epimeron 2, pereo-
podal dactyls, head and telson; but Bumeralius has
the typical antenna 1 of the urohaustorius form.
The first antenna resembles that in the Urothoe
group of genera. Prantinus differs from Urothoe,
Urothoides, and Carangolia in the absence of a
ventrolateral cephalic cheek, the uniarticulate
palp of maxilla 1 and the setose (rather than
spinose) rami of uropods 1-2 (albeit each bears
only 1 seta). Antenna 2 appears to be of more
characteristic haustorius form in Prantinus than in
those other taxa though antenna 2 of many spe-
cies of Urothoe is poorly known.
Prantinus appears to lie in a non-direct plesio-
morphic position from South African Cunicus,
though Prantinus possesses some of the strange
character of that taxon. Cunicus has somewhat
similar coxae, and the mandibular molar appears
to be reduced to a single spine. Cunicus, however,
has the sharp cephalic cheek and biarticulate
palp of maxilla 1 found in the urothoid group,
and furthermore, apomorphically has lost the
rami of uropods 1-2; has a poorly developed
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
inner plate of maxilla 1; weak dactyl of the
maxillipedal palp; and specializations in antenna
2, pereopod 5, and the hands of the gnathopods.
Prantinus talanggi, new species
Figures 13-18
DIAGNOSIS.?With the characters of the genus.
DESCRIPTION OF FEMALE.?Holotype female
"o," 7.19 mm: Head about 85% as long as wide,
rostrum about 16% as long as remainder of head
(rostrum measured only from final acclivity;
head, however, extended forward from antennal
notch), eyes represented by granular tissue
patches, no distinct ommatidia, ocular ganglia
visible. Facial formula of setae on article 1 of
antenna I = about 5 pairs of long to short setae
in irregular sets; on article 2, scattered; primary
flagellum with 6 articles, aesthetasc formula = 1-
3-3-1-1-0; accessory flagellum of 7 articles. Ar-
ticle 3 of antenna 2 with 6 large plumes; lateral
face of article 4 with mixture of short spines (21),
long plumose setae (10) and penicillate setae (3);
of article 5 with 5 plumes and 1 penicillate seta;
flagellum of 9 articles. Each mandibular incisor
with 3 teeth; right rakers 11, left 12 plus 1 hairlike
rudimentary; each molar with large main cusp
bearing long thin basal accessory cusp and ap-
posing minor cusp on each side more apical, plus
articulate whiplike seta from base opposite to
largest accessory cusp; article 3 of palp slightly
longer than article 2, latter with 3 inner apical
setae and 4 facial setae in tandem, spine formula
on right and left article 3 = 5-1-6-1. Inner plate
of maxilla 1 not attenuate, bearing 3 apicolateral
setae, 1 smaller seta directly at apex, pair of
mediofacial setae; outer plate with 10 spines;
palpar setae all apical. Inner plate of maxilliped
with 3 stout teeth, 2 medial and 2-3 apical setae;
outer plate with about 6 apical and lateral setae;
palp article 2 with 3 apicofacial setae, article 3
with pair of facial setae.
Coxa 1 broadly subrectangular, bearing 1 long
ventroposterior plume and several marginal se-
tules; coxa 2 deeper, with 7 long plumes and
several setules; coxa 3 almost tear-drop shaped,
with 14 plumes mostly posterior and several se-
tules; coxa 4 with about 17 ventral and 10 pos-
terior setae, long and plumose to short and poorly
plumose.
Setal and spine formulas on pereopod 3 = 6, 8,
2-1^1-1-1, 7 + 3 + 1; on pereopod 4 = 5, 7, 2-
1 ? 1 ?1?1, 7 + 3 + 1; margins of articles 5-6 not
serrate. Article 2 of pereopods 5-7 densely setose
posteriorly; dactyl of pereopod 5 with small an-
terior tooth.
Peduncular spine formulas of pleopods 1 -3 =
2 and 1, 2 and 0, 2 and 0; segmental formulas =
18-10, 13-6, 17-12 (inner proliferate); basal setal
formulas = 11-2-3-6, 1-1-1-1, 11-2-2-7.
Epimeron 1 sinuous posteroventrally; epimeron
2 extended posteroventrally, with 6 marginal pos-
teroventral setae, 1 facial seta near corner and
row of 3 facial setae more dorsally; epimeron 3
rounded-quadrate.
Apicolateral corner of peduncle on uropod 1
with 3 small spines (one hidden in illustration),
of uropod 2 with 4 spines; medial margin of each
with 7 and 5 setae in tandem respectively; dorsal
setae = 0 and 0, setae of rami = 1-1 and 1-1;
inner rami extending 87% and 83% along outer.
Apicolateral corner on uropod 3 peduncle with 6
plumes, medial margin with 4 plumes in 3 groups,
ventromedial apical with about 5 mixed setae,
spine formula on lateral margin of article 1 on
outer ramus of uropod 3 = 0, medium plumose
setae = 2-2.
Telson about 1.4 times as wide as long, weakly
alate laterally, cleft about 0.75 of its length, each
apex with 6 long to short plumes (almost penicil-
late in smaller members), each side with 2 small
penicillate setules.
Glands discernible only in pereopods 6-7.
Female tkq," 6.87 mm: Peduncles of uropods
1-2 with 2 and 1 dorsal setae.
DESCRIPTION OF MALE.?Male "m," 6.22
mm: Eyes large (illus.). Antenna 1 (illus.),
shorter and stouter than in female, article 2
broadened apically, article 3 densely furnished
with aesthetascs; formula on aesthetascs on pri-
mary flagellum = 0-1-3-1-1-0; accessory flagel-
NUMBER 360 39
Al
FIGURE 13.?Prantmus talanggi, new species, holotype, female "o," 7.19 mm (m = male "m."
6.22 mm; n = male "n," 6.02 mm; p = juvenile "p," 3.23 mm).
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
KGl
FIGURE 14.?Prantinus talanggi, new species, holotype, female "o," 7.19 mm (m = male "m,
6.22 mm; p = juvenile "p," 3.23 mm).
lum with 6 articles. Antenna 2 generally as in
female. Mouthparts, except for upper and lower
lips, distinctive, generally degenerative; right la-
cinia mobilis indistinct from rakers; left short,
deeply bifid; each palp article 3 spine formula =
4-1-5-1, each article 2 with 5 setae (illus.); max-
illae stunted (illus.); inner plate of maxilliped
stunted (illus.), outer plate not as well developed
as in female, facial setae on article 3 of palp
longer than in female.
Coxa 2 with 6 setae, coxa 3 with 12 setae
(elongate only); gnathopods as in female.
Armament formulas of pereopod 3 = 5, 7, 2-
1-1-1-1,6 + 3 + I;ofpereopod4 = 4, 5, 2 -1-1-
1,6 + 3 + 1. Pereopods 5-7 illustrated; article 2
of pereopod 5 with denser anterodorsal setulation
than in female; unlike female, article 2 of pereo-
pods 6-7 lacking almost all fully marginal seta-
tion, mediofacial row much better developed.
Pleopods illustrated, noting pleopod 2 better
developed than in female.
Epimera 1-2 better developed than in female
(illus.), epimeron 2 with numerous ventral and
facial setae.
Uropods 1-3 (illus.) more setose than in female;
peduncle of uropod 1 with 4 lateral and 2 dorsal
setae, apicolateral corner with only 1 spine and
setule; of uropod 2 with 1 dorsal seta and apico-
lateral armaments elongate and plumose; all setae
of peduncle on uropod 3 smaller than in female
but rami much more setose, inner ramus en-
larged, formula of short setae laterally on article
1 of outer ramus = 1-1-1-1-1-1, of long setae
= 1 ? 1 ? 1 ? 1 ? 1 ? 1 .
Telson (illus.) lacking alate appearance, ap-
pearing tumid, setae shorter than in female.
Male "n," 6.02 mm: Eyes poorly developed
(illus.). Flagella of antenna 1 with 5 and 6 articles.
Article 3 of antenna 2 with 5 plumes, armament
row on article 4 with 10 plumes, 13 spines, and 3
penicillates; article 5 with 7 plumes in arc, no
calceoli; flagellum of 7 articles. Right and left
rakers = 10 plus 2 rudimentaries and 9 plus 2
rudimentaries; palp article 2 with 4 setae, 2 of
these apical; setal formula on each palp = 3 -1 -
4-1. Maxillae, maxillipeds, and gnathopods like
female except maxilla 1 with 11 spines on 1 outer
plate, 10 on other. Coxae 1-3 illustrated. Spine
and setal formulas on pereopod 3 = 5, 7, 2 -1-1-
1-1, 7 + 3 + 1; on pereopod 4 = 4, 5, 2-1-1-1,
NUMBER 360 41
nE3
FIGURE 15.?Prantinus talanggi, new species, holotype, female "o," 7.19 mm (m = male "m."
6.22 mm; n = male "n," 6.02 mm).
P5t
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 16.?Prantinus talanggi, new species, holotype, female "o," 7.19 mm (m = male "m,
6.22 mm; n = male "n," 6.02 mm; p = juvenile "p,'? 3.23 mm; Y = pleopod; Z = gill).
NUMBER :5()() 43
FIGURE 17.?Pranlinus talanggi, new species, holotype, female "o." 7.19 mm (m = male "m."
6.22 mm;/? = juvenile "p." 3.23 mm, Z = gill).
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
D7
FIGURE 18.?Prantinus talanggi, new species, holotype, female "o," 7.19 mm (m = male "m,
6.22 mm; n = male "n," 6.02 mm; p = juvenile "p," 3.23 mm; Z = gill).
NUMBER 360
6 4- 3 + 1. Pereopods 5-7 like female. Pleopods
2-3 illustrated. Epimeron 2 (illus.) with 4 poster-
oventral and 4 facial setae. Apicolateral corner of
peduncles on uropods 1-3 with 2, 2, and 5 ele-
ments, like female in morphology; medial edges
of peduncles with 7, 5, and 2 elements; ventro-
medial setae of uropod 3 = 4; formula of short
setae on article 1 of outer ramus on uropod 3 =
1-1, long setae = 0-1. Telson (illus.) with only 3
apical setae on each lobe.
DESCRIPTION OF JUVENILE (juvenile "p," 3.23
mm).?Only ocular ganglia visible (illus.). Pri-
mary and accessory flagella of antenna 1 with 5
and 4 articles, formula of aesthetascs on primary
= 0-2-1-1-0. Armament row on article 4 of
antenna 2 with 7 plumes, 3 spines, and 2 penicil-
lates; article 5 with 3 plumes; article 3 with 3
plumes; flagellum with 5 articles.
Mandibular molar with only 3 cusps (illus.),
right lacinia mobilis deeply bifid (illus.); right
rakers 7 plus rudimentary; spine formula on each
palp article 3 = 2-1-3-0; palp article 2 with only
2 setae. Inner plate of maxilla 1 with 2 apical and
pair of medial setae only; outer plate with 9
spines. Inner plate of maxilla 1 with only 1 seta
in medial row. Following like adult female but
with fewer armaments: upper and lower lips,
maxilliped, gnathopods.
Coxa 2 with 2 long setae; coxa 3 with 4 long;
coxa 4 with 10 long setae; coxae 6-7 each with
only 2 long posterior setae.
Spine and setal formulas of pereopod 3 = 2, 2,
2-1, 3 + 1 + 1; of pereopod 4 = 2, 2, 2-1, 2 + 1
+ 1. Setae of pereopods 5-7 much fewer than in
adult. Epimeron 2 with 2 ventral and 1 facial
setae.
Peduncular spine formulas of pleopods 1-3 =
2 and 0, 2 and 0, 2 and 0; segmental formulas =
8-5, 6-3, 8-6; basal setal formulas = 8-1-1-3, 1-
1-1-1, 6-1-1-3; inner ramus of pleopod 2 lacking
basal clothespin spine.
Apicolateral corners of peduncles on uropods
1-2 naked; medial setae, 2 and 1 only. Uropod 3
(illus.) with article 2 on outer ramus elongate,
article 1 with only 1 lateral seta. Telson (illus.)
with only 1 long and 1 very short apical setae
tightly contiguous.
OBSERVATIONS.?Lateral penicillate setule pairs
on telson attached ventrally (not dorsally as in
other haustorioid taxa).
NSW specimens smaller than Victorian. Coxa
1 slightly more deeply emarginate ventrally; ar-
ticle 2 of pereopod 7 with posteromedial facial
ridge more pronounced.
ILLUSTRATIONS.?Antenna 1 of holotype drawn
with twist towards apex commencing at base;
maxilla 1 enlarged much more than maxilla 2,
whereas maxillae of male "m" drawn to same
magnification; gills poorly preserved, perhaps re-
constructed with slight inaccuracies. Uropods 1-
2 of "n" projecting laterally and dorsally as in
most specimens, not flattened or altered in this
view.
HOLOTYPE.?NMV, female "o," 7.19 mm (il-
lus.).
TYPE-LOCALITY.?CPBS C6/4, Western Port,
Victoria, 30 Mar 1964, 10.2 m, fine sand.
VOUCHER MATERIALS.?Type-locality: male
"n," 6.02 mm (illus.), juvenile "p," 3.23 mm
(illus.); CPBS A3/5: female "q," 6.87 mm; CPBS
25N/3: male "m," 6.22 mm (illus.); AM P.23327
OHDWBS, off Newcastle, NSW, transect
06.03.03.04: female "t," 5.43 mm.
MATERIAL.?CPBS, 3 samples from 3 stations
(10); WPBES 1 sample (1); AM HDWBS 4 sam-
ples (4).
DISTRIBUTION.?Victoria, Western Port; NSW,
offshore from Burwood Beach, 16 km south of
Hunter River; 10-28 m, sand.
UROHAUSTORIIDAE, new family
DIAGNOSIS.?Rostrum weak, head short, cheek
poorly developed. Antenna 1 of haustorius form,
article 1 short, articles 2-3 progressively shorter,
weakly geniculate or not, flagella elongate, usu-
ally subequal to peduncle. Antenna 2 of hausto-
rius form, article 4 expanded, article 5 shorter
and narrower than article 4, these articles fur-
nished with 1 or more longitudinal rows of facial
armaments, ventral margin of article 4 with at
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
least 3 kinds of setae: (1) elongate plumes, (2)
shorter and suffer glassy spines set in clusters or
ranks, and (3) usually bulbar-based penicillate
setules; flagellum subequal to or longer than ar-
ticle 4 of peduncle. Prebuccal complex massive,
upper lip usually dominant and epistome scarcely
distinct. Mandibles bearing elongate, poorly
toothed incisors; laciniae mobiles present on each
side, diverse; rakers distinct, usually serrate, but
few in number; molar large and weakly tritura-
tive, usually furnished with accessory chopper;
palp 3-articulate, article 3 with numerous outer
setae, most setae awned. Lower lip with discrete
inner lobes, mandibular extensions of outer lobes
well developed. Maxilla 1 with uniarticulate palp,
inner plate with fewer than 5 setae. Maxilla 2
ordinary7, inner plate with oblique but poorly
developed facial row of setae. Maxillipeds with
unexpanded bases, normally enlarged plates,
outer spinose; palp 4-articulate, article 2 ex-
panded, article 4 clavate, multisetose. Maxillae
and maxillipeds lacking baler lobes.
Coxa 2 very small, not forming stepped inter-
grade between coxae 1 and 3; coxa 3 dominant
or not dominated by coxa 4, broadly extended
posteroventrally. Coxal gills on segments 2-5.
Brood plates slender.
Gnathopods feeble, grossly alike in proportions,
wrists elongate, but gnathopod 1 simple, gnath-
opod 2 subchelate or minutely parachelate; arti-
cle 3 short. Article 5 of pereopods 3-4 broad,
slightly expanded, not deeply lobate, with thick
posterior spines; dactyls of pereopods 3-5 well
developed, those of pereopods 6-7 variable; per-
eopod 5 of haustorius form, articles 2, 4, and 5
expanded, articles 5 and 6 with extensive facial
rows of spines; pereopods 6-7 alike, articles 5-6
weakly expanded; no pereopod with underslung
articulation.
Pleopod 2 usually inferior in size, number of
articles or setation; peduncles of pleopods not
longer than wide, coupling hooks paired; inner
rami inferior, each usually bearing 1 clothespin
spine.
Epimeron 1 scarcely developed; epimeron 2
dominant in setation, often dominant in size.
Urosomites ordinary, though often furnished with
lateral teeth. Rami of uropods 1-2 linguiform,
setose (not spinose); uropod 3 of ordinary haus-
torioid-phoxocephalid kind, outer ramus domi-
nant, 2-articulate, peduncle short, flat, expanded,
rami poorly setose apically.
Telson variable. Sexual dimorphism weak.
TYPE GENUS. ? Urohaustonus Sheard.
COMPOSITION.?Gheegerus, new genus; Narumus,
new genus; Tuldarus, new genus; Tottungus, new
genus; Dirimus, new genus.
Relationship.?This family differs from the
Urothoidae in the poor development of the ven-
tral cephalic cheek, the full development of the
haustorius antenna 2, and in the dominance of
epimeron 2. All taxa have linguiform and setose
rami on uropods 1-2 whereas urothoids have
styliform and spinose rami. The urothoid Cumcus
upsets easy recognition and clear distinction be-
tween the 2 groups because the rami of uropods
1-2 are absent and the peduncles are linguiform
and setose.
Distinguishing characters of Haustoriidae are
listed with the diagnosis of that family below.
Phoxocephalopsidae bear styliform or rod-
shaped spinose rami on uropods 1-2.
Urohaustoriidae differ from Zobrachoidae in
the simple gnathopod 1; the great reduction in
size of coxa 2 (and, in all but one case, of coxa 1);
enlargement and posteroventral extension of coxa
3; almost full loss of epimeron 1; to a lesser degree
the uniformly small rostrum or simple outline
dorsally of the head; the small number of setae
on the palp of maxilla 1; reduction in the exten-
sion and cuspidation of the mandibular molar;
and generally the reduction in size of article 2 on
the outer ramus of uropod 3.
Prantinns of the Zobrachoidae intergrades some
of these characters slightly. Epimeron 1, for ex-
ample, is not as well developed as in other zobra-
choids; article 2 on the outer ramus of uropod 3
is not as greatly elongate; but on the other hand
the mandibular molar is extremely cuspidate and
the head is strongly rostrate or, at least, extended
forward dorsally.
NUMBER 360 47
Key to Genera of Urohaustoriidae
Dactyl of pereopod 5 spinose 2
Dactyl of pereopod 5 not spinose 4
Antennae bearing supernumerary oar-shaped setae, article 2 of pereopods
6-7 narrow or strongly tapering distally Narunius, new genus
Antennae lacking supernumerary oar-shaped setae, article 2 of pereopods
6-7 fully expanded, not tapering distally 3
Coxa 3 large, coxa 4 small Urohaustorius
Coxa 3 small, coxa 4 large Gheegerus, new genus
Coxae 1 and 2 of equal size, right and left laciniae mobiles of subequal
size, thin, epimeron 2 about equal in size to epimeron 3, lacking
setae Dirimus, new genus
Coxa 1 larger than coxa 2, left lacinia mobilis much broader than right,
epimeron 2 either larger than or more setose than epimeron 3 5
Gnathopod 2 subchelate, dactyls of pereopods 5-7 setose in
female Tottungus, new genus
Gnathopod 2 parachelate, dactyls of pereopods 5-7 not setose in
female Tuldarus, new genus
Urohaustorius Sheard
['rohaustonus Sheard, 1936:445-446.
DIAGNOSIS.?Rostrum weak. Peduncle of an-
tenna 1 short, stout, articles 2 and 3 of peduncle
progressively shortened, weakly geniculate, both
flagella elongate. Antenna 2 of haustorius form,
article 4 expanded, article 5 small, articles 4-5
with facial armaments, article 4 with long ventral
setae and subventral clusters of bifid setae. Aes-
thetascs linear. Mandibular incisors extended,
thin, essentially untoothed; right and left laciniae
mobiles not alike; rakers usually serrate, 2 on
right mandible, 3 on left; molar broad and bulky,
not strongly extended, with weak triturative sur-
face and cuspidate accessory chopper; setae of
palp article 3 awned. Mandibular lobes of lower
lip well developed. Inner plate of maxilla 1 small,
slender, poorly setose; outer plate with 11 spines;
palp slightly shortened, uniarticulate, palpar se-
tae attached in 2 sets. Inner plate of maxilla 2
with weakly submarginal oblique row of setae.
Outer plate of maxilliped with spines; palp article
2 expanded, article 3 elongate, dactyl clavate,
multisetose. Maxillae and maxillipeds lacking
baler lobes.
Coxae 1-4 not progressively larger, coxae 1-2
very small, usually equal in size, most of coxa 2
hidden by coxa 3, latter much larger than coxa 4
and hatchet-shaped, extended posteroventrally;
coxae 2-5 each with simple gill; brood plates 4
pairs, slender.
Gnathopods small, dissimilar, gnathopod 1
simple, gnathopod 2 weakly chelate (parache-
late), wrists elongate, hand of gnathopod 2 weakly
mitelliform. Dactyls of pereopods 3-7 distinguish-
able; those of pereopods 3-5 large, of pereopods
6-7 variable, large to vestigial; dactyl of pereopod
5 large and spinose. Article 2 of pereopods 5-7
expanded; pereopod 5 of haustorius form; distal
articles of pereopods 6-7 not underslung, only
moderately widened, pereopods 6-7 otherwise
similar, weakly dominating pereopod 5.
Pleopod 2 inferior, peduncles of pleopods
short, wider than long, inner rami moderately
shorter than outer. Epimeron 1 scarcely differen-
tiated, epimeron 2 dominantly setose. Urosomite
1 scarcely produced and poorly setose ventrally.
Rami of uropods 1-2 linguiform, though inner
rami often reduced or vestigial, marginally setose;
peduncle of uropod I strongly setose laterally; of
uropod 2 spinose or poorly setose laterally. Uro-
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pod 3 of ordinary form, outer ramus dominant
and biarticulate, article 2 with 2 large apical
setae. Telson short, broader than long, or not
greatly longer than broad, cleft to entire.
DESCRIPTION.?Eyes indistinct or very poorly
discernible, ocular ganglia visible. Dorsolateral
surface of article 1 on antenna 1 furnished with
longitudinal row of armaments; article 2 densely
setose dorsolaterally, groups distinguishable, ar-
ticle 3 poorly setose. Article 3 of antenna 2 short,
poorly setose, flagellum much longer than article
4 of peduncle.
Right lacinia mobilis linguiform; left broader
and usually bifid; article 3 of mandibular palp
shorter than article 2. Lower lip usually with 1
cone on each outer lobe. Inner plate of maxilla 1
with very few short apical setae and one large
apicomedial seta; spines on outer plate 11, mostly
bifid. Inner plate of maxilliped with 2 stout apical
spines (a third false spine marked by plumes).
Coxae 1-2 usually poorly setose. Gill 2 dominant,
gill 5 rarely larger, often much smaller than
posterior lobe of coxa 5. Gnathopod 2 bearing
surficial button combs.
Pereopods 3-4 lacking extensive secondary fa-
cial rows of spines on article 5.
Uropods 1-2 with huge dorsal setae on pedun-
cles, rami lacking basoventral setae, medial mar-
gins of peduncles usually heavily setose.
TYPE-SPECIES.?Urohaustonus halei Sheard, 1936
(original designation).
COMPOSITION.?Urohaustonus vercoi Sheard, U.
metungi Fearn-Wannan, and the following new
species: U. pulcus, U. pentinus, U. merkanius, U.
wingaro, U. parnggius, U. yurrus, U. urungari, U.
perkeus, and U. gunni.
RELATIONSHIP.?Apart from the uniarticulate
palp of maxilla 1, Urohaustonus retains relatively
ordinary mouthparts but is 1 of the most special-
ized taxa in terms of coxae. The first 2 coxae are
very small (except for U. gunni, with enlarged
coxa 1), but coxa 3 is the largest of any and is
strongly extended posteroventrally. Rami of uro-
pods 1-2 are linguiform and setose (apart from
several species with reduced inner rami). Gnath-
opod 1 is simple as in the Haustoriidae.
Urohaustonus may bear closest superficial resem-
blance to the Haustoriidae, but differs in the
uniarticulate palp of maxilla I, retention of man-
dibular lobes on the lower lip, the extended and
toothed mandibular incisors, retention of the
maxillipedal dactyls, greatly reduced setation on
the inner plate of maxilla 1, presence of an orga-
nized dorsolateral row of armaments on article 1
of antenna 1, the longer accessory flagellum, ab-
sence of a secondary facial row of armaments on
article 4 of antenna 2, strongly reduced coxae 1-
2 and dominant coxa 3, the unlobate article 5 of
pereopods 3-4 bearing dominant posterior spi-
nation and no facial armaments, the strongly
discernible dactyls of pereopods 3-5, the close
similarity of pereopods 6-7 between themselves,
dominance of epimeron 2, the fully linguiform
rami of uropod 1, presence of long lateral setae
on the peduncle of uropod 1 (contrasted to short
spines), lack of dominance in article 2 on the
outer ramus of uropod 3 and reduction of apical
setation on that article to a set of 2 elongate setae,
other normal shapes and setation of uropod 3,
and the general appearance of the telson, which
in Haustorius has a gaping cleft and manifold
basolateral setal brushes.
The loss of prehensility of gnathopod 1 and
dominance of coxa 3 distinguish Urohaustonus
from the otherwise similar Zobracho, Bumeralius,
and Prantinus.
Urohaustonus differs from Narunius primarily in
the true dominance by coxa 3.
IDENTIFICATION OF CRYPTIC SPECIES.?A gen-
eral description section is attached to U. merkanius,
U. wingaro, U. parnggius, U. yurrus, and U. urungari
because these were presumed to be cryptic sibling
species of greater than normal difficulty in rec-
ognition. We now believe that 5 species are in-
volved but that they are incompletely segregated
by morphological characters and differ from each
other in different habitats by varying degrees of
character expression.
It is impossible to identify many of these juve-
niles satisfactorily.
THE pulcus COMPLEX OF UROHAUSTORIUS.?At
least 9 major morphs occur in this collection of
NUMBER 360 49
specimens from diverse sources. These morphs are
U. pulcus, U. pulcus morph 872, U. pentinus, U.
merkamus, U. wingaro, U. parnggius, U. yurrus, U.
urungan, and U. species M. We believe the com-
plex is morphologically too diverse to embody a
single species, even though it has been found
impossible to make perfect typological groupings
within it. Because over 95% of the specimens do
fall into definable groups (which we assume to be
incipient species), however, we have called most
of the morphs separate species as a practical
recourse for ecologists seeking identifications in a
gross survey. Perfect typological groupings would
require a very large number of species to be
separated from the specimen pool, but coales-
cence of variants into a few species requires
acceptance of several unusual replicates
("mutants") that would have to be assigned to
different species, depending on characters other
than the mutant. For example, the presence of
subsidiary setules on the dactyl of pereopod 5
would require such specimens to be considered as
"mutant" (phenotypic) forms occurring in 2 dif-
ferent species; or the ventral displacement of
basal elements in the main armament row on
article 4 of antenna 2 would require allocation of
such specimens to two different species.
More than 95% of our adult specimens could
be divided into 3 species based on degree of
telsonic cleavage; deeply, moderately, or not cleft.
But this result is clearly unsatisfactory as speci-
mens with uncleft telson are divisible into those
with or without double spination on the dactyl of
pereopod 5. If search for subdivisions were to stop
at this point, four reasonably well-defined species
could be proposed (pulcus, pentinus, merkamus,
parnggius). Two other kinds of clear alternative
character expressions do occur: (1) ventral dis-
placement of elements in the main armament row
on article 4 of antenna 2; and (2) presence or
absence of accessory setules adjacent to the spines
on the dactyl of pereopod 5 in specimens other-
wise having only single spines in tandem. Char-
acter 2 represents a perfect intermediate between
specimens with single spines and doubled spines
on the dactyl of pereopod 5 because there is no
intermediacy in size of the setules (see female "t"
of morph 872, an exception, page 60.) No inter-
mediates between even and displaced armaments
on antenna 2 have been found.
When specimens with displaced armament on
antenna 2 (U. wingaro and U. parnggius) are pooled
together, the degree of telsonic cleavage is found
to be more than slightly variable; but when spec-
imens with accessory setules on dactyl 5 are
pooled (morph 872) a much larger variability in
telsonic cleavage is found in the aggregate, the
degree extending from well cleft to poorly cleft.
This would suggest that either (1) degree of cleft-
ing is a poor specific character (variable), or (2)
that presence of accessory setules is a replicative
(mutant or recombinant) character in 2 species.
But degree of cleavage (already) accounts for 95%
accuracy in classifying specimens and is a major
character of value in other species groups of the
genus and other genera of the family group.
More divisions are possible. Specimens with 2
long proximal plumose setae in the armament
row on article 1 of antenna 1 (morph M) are rare
and mostly confined to 1 sample but we have
found this condition also in a specimen of U.
metungi outside this specific complex and also in
2 probable U. pulcus juveniles.
Several of the so-called species have correlative
or associative characters confirming more or less
that each is a definitive species; these characters
are discussed in their respective places.
Couplets 3-12 of Key I to Urohaustonus are
written in such a way that U. pulcus is situated
last and represents the primary name to be ap-
plied to this complex. Taxonomists wanting to
divide the complex into its parts can decide where
in other couplets to make the division: for exam-
ple, couplets 5-6 can be ignored but the speci-
mens remain divisible into species in couplets 7-
12; or couplets 7-8 can be ignored and the spec-
imens still remain divisible into grosser species in
couplets 9-12; couplet 9 could then be ignored
and the specimens would be divisible into 2 spe-
cies; or couplet 10 could be ignored and the
specimens called U. pulcus.
We interpret the material to represent 7 good
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
species (pulcus, pentinus, rnerkamus, parnggius, win- in spine doubling on pereopod 5, but to some
garo, yurrus, and urungari), and 1 potential species degree these geographic clines are controlled by
(species M), plus a replicate morph, (872), assign- sediment conditions and not by temperature. We
able to pulcus; but we recognize that few juveniles n o t e t h a t appropriate Queensland sediments are
and onlv 95% of the adults can be identified probably coarser than Victorian sediments so that
.
 r ' ., ^ n ir an increase in value for doubled spines and larger
satisfactonlv. Gene flow is therefore apparent , r , , ? ,
, . , ? , , bodies would occur in Queensland, which goes
among the species, represented either by com- . . . ^ c , ,s
 . , , ? , ? ? ? counter to thermal clines. h>uch counter selection
monlv occurring hvbridization or bv genetic re- ,
 r , f ? r
-
 5
 - -
 &
 may therefore make for a very complex specific
combinations occurring frequently, themselves moroholoev
reminiscent of earlier hybridization. Gene flow is
 W e h a v e c o n s t r u c t c d 3 different keys to the
sufficiently sluggish and adequately selected species of the genus so that an identifier can have
against by environmental benefits to certain 3 different starting places. Specimens that fail the
morphs in a degree permitting maintenance of test 2 or 3 times must be considered as possible
ecophenotypes, if not ecogenotypes. There is also undescribed taxa, but study of descriptions may
weak evidence that clines may occur, for example, demonstrate close affinities.
Key I to Selected Species of Adult Urohaustorius
(('. vercoi omitted)
1. Rami of uropods 1-2 subequally extended 2
Inner rami of uropods 1-2 much shorter than outer rami 4
2. Coxa I much larger than coxa 2, forming long anterior curved hook,
coxa 7 with long posterior spike 17. gunni, new species
Coxa 1 not larger than coxa 2, coxa 7 lacking spike 3
3. Telson with simple marginal setae in addition to penicillate setules,
posterior setae on article 4 of pereopod 6 not continuous to apex, small
gap occurring between ultimate setae and spine at corner, dactyl of
pereopod 5 with some spines in pairs U. halei
Telson without simple marginal setae, otherwise only with 6 penicil-
late setules, posterior setae on article 4 of pereopod 6 continuous
to apex, no small gap occurring between ultimate seta and spine
at corner, dactyl of pereopod 5 with all spines set singly in tandem
U. metungi
4. Dactyls of pereopods 6-7 vestigial, coxae 1-2 rectangular and well setose
U. perkeus, new species
Dactyls of pereopods 6-7 well developed, coxae 1-2 triangular or trape-
zoidal, poorly setulose 5
5. Dactyl of pereopod 5 with some spines or elements set in pairs 6
Dactyl of pereopod 5 with all spines set singly in tandem 7
6. Dactyl of pereopod 5 with at least one spine position paired by second
spine U. merkanius, new species
Dactyl of pereopod 5 with at least one spine position paired by accessory
setule V. pulcus, morph 872
7. Proximal part of main armament row on article 4 of antenna 2 displaced
ventrally g
Armament row on article 4 of antenna 2 evenly oblique 9
NUMBER 360 51
8. Tclson weakly cleft U. wingaro, new species
Telson entire L7. parnggius, new species
9. Armament row on article 1 of antenna 1 with 2 long setae proximally;
article 2 of pereopod 6 narrow U. species M
Armament row on article 1 of antenna 1 with 0-1 simple seta, article 2
of pereopod 6 broad 10
10. Telson entire, glandular (male sternite 6 with blunt transverse
process(es)) 11
Telson weakly to strongly cleft, not glandular (male sternite 6 with sharp
axial process) 12
11. Telson of tall extended form, (male sternite 6 with central broad ridge,
bilaterally humped) U. urungari, new species
Telson of broad unextended form, (male sternite 6 with central mound
only) U. yurrus, new species
12. Telson deeply cleft, article 2 of pereopod 7 with proximoposterior setae
U. pulcus, new species
Telson weakly cleft, emarginate, article 2 of pereopod 7 with stout
proximoposterior spines U. pentinus, new species
Key II to Selected Species of Adult Urohaustorius
(U. vercoi and V. pulcus, morph 872, omitted)
1. Armament row on article 1 of antenna 1 with some stout spines .... 2
Armament row on article 1 of antenna 1 with fine setae or medium
spines only 4
2. Cleft of telson forming deep split U. pulcus, new species
Cleft of telson obsolescent or absent 3
3. Telson excavate U. pentinus, new species
Telson entire 9
4. Basal pair of elements in armament row on article 1 of antenna 1
plumose U. species M
Basal pair of elements in armament row on article 1 of antenna 1 simple
5
5. Basal part of armament row on article 4 of antenna 2 evenly oblique
' 6
Basal part of armament row on article 4 of antenna 2 displaced ventrad
10
6. Armament row on article 4 of antenna 2 mixing spines and setae
U. merkanius, new species
Armament row on article 4 of antenna 2 consisting of setae only 7
7. Telson cleft, setae of coxa 1 dense, inner rami of uropods 1 and 2
significantly shortened L7. perkeus, new species
Telson entire, setae on coxa 1 sparse, inner rami of uropods 1 and 2
extending subequally 8
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
8. Coxa 1 sickle shaped U. gunni, new species
Coxa 1 ordinary 11
9. Telson of tall extended form, (male sternite 6 with central broad ridge,
bilaterally humped) U. urungari, new species
Telson of broad unextended form, (male sternite 6 with central mound
only) U. yurrus, new species
10. Telson weakly cleft U. wingaro, new species
Telson entire U. parnggius, new species
11. Telson with simple marginal setae in addition to penicillate setules,
posterior setae on article 4 of pereopod 6 not continuous to apex, small
gap occurring between ultimate setae and spine at corner, dactyl of
pereopod 5 with some spines in pairs V. halei
Telson without simple marginal setae, otherwise only with 6 peni-
cillate setules, posterior setae on article 4 of pereopod 6 continuous
to apex, no small gap occurring between ultimate seta and spine
at corner, dactyl of pereopod 5 with all spines set singly in tandem
U. metungi
Key III to Selected Species of Adult Urohaustorius
(U. vercoi and U. pulcus, morph 872, omitted)
1. Telson cleft or emarginate 2
Telson entire 5
2. Coxa 1 with long setae, tooth of epimera 1-2 large, dactyls of pereopods
6-7 vestigial 17. perkeus, new species
Coxa 1 with small setules, tooth of epimera 1-2 small or absent, dactyls
of pereopods 6-7 ordinary 3
3. Proximal part of main armament row on article 4 of antenna 2 displaced
ventrad, male sternite 6 with blunt process
U. wingaro, new species
Armament row on article 4 of antenna 2 even, oblique, male sternite 6
with sharp tooth 4
4. Cleft of telson deep, article 2 of pereopod 7 with long proximoposterior
setae U. pulcus, new species
Cleft of telson obsolescent, article 2 of pereopod 7 with proximoposterior
stout spines U. pentinus, new species
5. Rami of uropods 1-2 extending subequally, apicolateral armament on
peduncle of uropod 3 of thin elements 6
Inner ramus of uropods 1-2 shortened strongly, apicolateral armament
on peduncle of uropod 3 of thick elements 8
6. Coxa 1 forming long curved spike U. gunni, new species
Coxa 1 rectangular, short 7
7. Article 4 of pereopod 6 with gap in posterior armaments, telson with
both penicillate and simple setules U. halei
Article 4 of pereopod 6 lacking gap in posterior armaments, telson with
only penicillate setules \J, metungi
NUMBER 360 53
8. Proximal part of main armament row on article 4 of antenna 2 displaced
ventrad, setae on posterior margin of article 2, pereopod 7 widely
spread U. parnggius, new species
Armament row of article 4 of antenna 2 evenly oblique 9
9. Article 2 of pereopod 6 narrow, first 2 setae in armament row of article
1 on antenna 1 plumose U. species M
Article 2 of pereopod 6 broad, first 2 setae in armament row of article 1
on antenna 1 simple 10
10. Epimera 2-3 with medium posteroventral tooth, male sternite 6 with
sharp ventral spike U. merkanius, new species
Tooth of epimera 2-3 obsolescent or absent, process on male sternite 6
blunt 11
11. Telson short and broad, spines predominating in main armament row
on article 4 of antenna 2, male sternite 6 process median, blunt
U. yurrus, new species
Telson tall, setae predominating in main armament row on article 4
of antenna 2, male sternite 6 process transverse, bilaterally humped
U. urungari, new species
Urohaustorius pulcus, new species
FIGURES 19-23
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 mixing spines and setae. Armament
row on article 4 of antenna 2 even. Accessory
flagellum reduced. Main flagellar articles of male
antenna 1 thickened, aesthetascs 2-many per ar-
ticle and present on article 1, article 5 of male
not bulbous. Coxa 1 not larger than coxa 2, both
bearing only tiny setules. Spines on dactyl of
pereopod 5 set singly in tandem. Dactyls of per-
eopods 6-7 well developed. Article 2 of pereopod
7 with 3 stout posterior setae and weak scattered
setules. Male pereonal sternite 6 with curved,
apically acute, posteriorly directed ventral tooth.
Epimeron 2 bearing small tooth and 3-8 facial
setae in horizontal row. Tooth of epimeron 3
obsolescent. Inner rami of uropods 1-2 much
shorter than outer rami. Dorsolateral margin of
peduncle on uropod 2 with stout spines. Apico-
lateral corner of peduncle on uropod 3 with stout
spines, outer ramus with setae and spines laterally
on article I. Telson distinctly cleft, forming slit,
bearing 2-3 apical setae on each half.
DESCRIPTION OF FEMALE.?Holotype female
"a," 3.72 mm: Head about 70% as long as wide,
rostrum about 14% as long as remainder of head.
Primary flagellum of antenna 1 with 10 articles,
I aesthetasc (rarely 2) on articles 4-9; accessory
flagellum 8-articulate; facial formula of article 1
on antenna 1 = E, 4S, E, 3S, 2P. Article 3 of
antenna 2 with 3 facial setae; main armament
row on article 4 in 2 weakly divided sections of
II and 18, spines dominant; on article 5 = 24;
article 5 = 45% as wide as long, about 70% as
long as article 4; flagellum of 12 articles. Right
incisor simple, left with 2 teeth, choppers on
molars with 4 cusps; formula on right palp article
3 = 9-1-3-1, (left aberrant, see illustration). Inner
plate of maxilla 1 with 1 short apical seta, 1 large
apicomedial seta; outer plate with 11 spines (only
9 shown in illustration). Inner plate of maxilla 2
with medial setae. Inner plate of maxilliped with
7 medial setae, outer plate with 1 apicolateral
seta, palp article 2 with 7 setae in facial row,
article 3 with 2 setae in facial row.
Coxa 1 broadly subtriangular; coxa 2 similar
but apex blunter; coxa 3 with 7 ventral setae all
in posterior half, posterior margin with 2 setae
both ventrad; coxa 4 with 7 ventral and 8 poste-
rior setae.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 19.?Urohaustonus pulcus, new species, holotype, female "a," 3.72 mm
3.97 mm; m = male "m," 3.60 mm).
= male "g,
NUMBER 360 55
mZP5 Mtv 0X1
FIGURE 20.? Urohauslonus pulcus, new species, holotype, female "a," 3.72 mm (c = female "c,"
3.62 mm; g ? male "g," 3.97 mm; m = male "m," 3.60 mm; V = pleopod, L = gill).
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
uKPS
* QGlt P5t
FIGURE 21.?Urohaustonus pulcus, new species, holotype, female "a" 3.72 mm (c = female "c,
3.62 mm; m = male "m," 3.60 mm; u = female "u," size unknown; Y = pleopod).
NUMBER 360 57
gW
FIGURE 22.?Urohaustonus putcus, new species, holotype, female "a," 3.72 mm (g = male "g,
3.97 mm; m = male "m," 3.60 mm; q = female "q," 3.48 mm; Y = pleopod).
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
R2
FIGURE 23.? Upper: Urohaustonus pulcus, new species, holotype, female "a," 3.72 mm (g = male
"g," 3.97 mm; a = male "a," 2.98 mm, morph 872 (=aberrant). Lower: Urohaustonus pentinus,
new species, holotype, female "w," 2.28 mm (n = male "n," 2.54 mm; x = female "x," 2.06 mm;
y = female "y," 2.08 mm; Y = pleopod).
Setal and spine formulas on pereopod 3 = 5, 5,
2 - l - U l - l , 6 + 5 + 1 ; pereopod 4 = 4, 5, 2 -1-1-
1-1, 6 + 4 + I; posterior margin of articles 5-6
with partially doubled row of mostly large even
serrations. Article 2 of pereopods 5-7 progres-
sively with medium to sparse setal densities pos-
teriorly, article 2 of pereopod 5 lacking mediofa-
cial brush, dactyl with 5 spines in tandem.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental forumulas
= 17-11, 14-10, 14-12 (inner proliferate); basal
setal formulas = 7-2-2-4, 2-1-1-1, 6-2-1-3.
Epimeron 2 with 4 setae in horizontal facial
Lateral margin of peduncle on uropod 1 with
7 setae, apicolateral corner with 1 setule; of uro-
pod 2 with 1 disjunct lateral spine and pair of
apicolateral spines plus setule, dorsal setae = 3
and 3; setae of rami = 8-3 and 8-3, inner rami
extending 60% and 47% along outer. Apicolateral
corner of peduncle on uropod 3 with 5 spines,
medial margin with 6 weak setae in tandem,
ventromedial apical row(s) with about 8 mixed
setae, spine formula on lateral margin of article
1 on outer ramus = 1-1-1-1, setal formula = 1-
1-1-1.
Telson about 0.87 as wide as long, cleft about
0.30 of its length, setal formula = 2P, 1, P, 2.
NUMBER 360 59
Lateral gland clusters well developed in pleon-
ites 3-6, more weakly developed in peduncle of
uropod 3.
Female "q," 3.48 mm: Spine formulas on
article 6 of pereopod 3 = 5 + 5 + 1; of pereopod
4 = 5 + 4 + 1; telson illustrated, formula same as
holotype but appearance distinctive.
DESCRIPTION OF MALE.?Male "m," 3.60
mm: Like female except in following propor-
tions and counts: Primary flagellum of antenna
1 thicker; article 4 of antenna 2 slightly narrower;
flagellum of antenna 2 with 10 articles; both
choppers of mandibular molars with 5 cusps;
right palp article 3 formula = 9-1-3-1, left = 8-
1-3-1; inner plate of maxilla 2 with 6 setae in
oblique row, outer plate with 2 apicolateral small
setae outside enlarged seta; spine formula of ar-
ticle 6 on pereopod 3 = 5 + 5 + 1; on pereopod
4 = 5 + 4 + 1, article 5 on pereopod 4 with 4
anterodistal setae; gill of coxa 5 scarcely larger
than posterior lobe; inner ramus of uropod 1 with
4 setae; ventromedial apex of peduncle on uropod
3 with 11 setae. See also illustrations of antenna
I, coxa 1, coxa 4, telson (formula = 3, P, (1-2),
2P).
Male "w," 3.56 mm: Apicolateral corner of
peduncle on uropod 3 with 4 spines and setule.
Primary flagellum with 13 articles, proximal 5
thickened; aesthetasc formula = 4-5-5-5-5-5-5-
5-5-5-2-2-0. (Only long aesthetascs are included
in the formula; the terminal 1 (0) is rudimentary.)
Article 5 of antenna 2 about 74% as long as article
4, about 54% as wide as long.
OBSERVATIONS.?Although the most common
number of spines on outer plate of maxilla 1 is
II, some specimens examined for this character
have 10 on 1 side and 11 on the other.
HOLOTYPE.?NMV, female "a," 3.72 mm (il-
lus.).
TYPE-LOCALITY.?CPBS A6/2B, Western Port,
Victoria, 28 Aug 1964, 10.6 m, sand and weed.
VOUCHER MATERIAL.?Type-locality: female
"c," 3.62 mm (illus.); female "u," 3.53 mm; male
"g," 3.97 mm (illus.); male "m," 3.60 mm (illus.);
CPBS 34S/1272: female "q," 3.48 mm (illus.);
CPBS24N/1: male "w," 3.56 mm.
RELATIONSHIP.?Although this is not the type-
species of the genus, it is used as the model
plesiomorph because of its deeply cleft telson
combined with easily visible dactyls of pereopods
6-7, unpaired spines on the dactyl of pereopod 5,
mixture of spines and setae in the facial rows on
antennae 1-2, and the absence of any other
"distinctive" features, such as huge teeth on epi-
mera or coxae. Because the rami are subequal,
the type-species, U. halei, has much more primi-
tive uropods 1-2, but U. halei also has doubled
spination on the dactyl of pereopod 5, somewhat
bizarre epimera, and an uncleft telson.
MATERIAL.?CPBS, 36 samples from 17 sta-
tions (148); WPBES, 4 samples from 4 stations
(9); PPBES, 5 samples from 4 stations (15); EBS,
1 sample (4).
DISTRIBUTION.?Victoria, Port Phillip Bay to
Port Macquarie, NSW, 2.8-16 m, sand, and
gravel, sand and mud, sand and weed.
Urohaustorius pulcus, morph 872
FIGURES 23 (part), 27 (part)
DIAGNOSIS.?Like U. pulcus, but dactyl of pe-
reopod 5 with 1 or more setules paired with single
spines in tandem; article 2 of pereopod 7 with 1
strong spine-seta posteroproximally and 1 weaker
seta near midmargin; telson generally cleft Vs,
with gaping lobes, but occasionally specimen with
telson cleft about Vs; some males (presumably
adult) with more than 2 aesthetascs per article on
main flagellum of antenna 1; usually 1 telsonic
seta almost perfectly apical on each lobe.
REMARKS.?Owing to the inconsistency of the
material, these specimens are difficult to treat as
a species. Unlike U. merkanius, in which consis-
tency of spine-doubling is high on the dactyl of
pereopod 5, a large percentage of morph 872
specimens shows the identificatory character only
on 1 side or the other. The character may appear
only when a damaged or broken dactyl regener-
ates. However, it has appeared associated with
uncleft telson only once, suggesting it is not a
regenerative character. Certain specimens of
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
morph 872, resembling U. pentinus in the poorly
cleft telson, cannot be assigned to that species on
account of the presence of setae (not stout spines)
on article 2 of pereopod 7, and the narrower outer
ramal articles of uropod 3. They might be hybrids
between U. merkamus and U. pulcus, in which the
second spine of a pair is represented only by a
setule but in which the telsonic expression is
variable or a mixture of conditions between par-
ents.
The evenness of the armament row on article
4 of antenna 2, the posterior margin of article 2
of pereopod 7, and the shape of uropod 3 separate
morph 872 from U. wingaro, and also from U.
parnggius. If they are hybrids, one of these two
might be a parent.
Morph 872 differs from most specimens of U.
pulcus in the posterior setation on article 2 of
pereopod 7 but a few specimens of U. pulcus have
been found with a similar formula. The gaping
telson of morph 872 is matched in large males of
U. pulcus bearing well-broadened articles of the
main flagellum. The real distinction from U.
pulcus lies in the presence of spines and setae on
the dactyl of pereopod 5.
The single specimen (female "1"), with uncleft
telson bearing setules on the dactyl of pereopod
5, is an apparent anomaly as it is the only such
specimen in 200 of the U. merkanius complex.
Morph 872 has been recovered from CPBS sta-
tions 24N (in samples taken years apart, in 1965
and 1972), 24S and 26 N, as well as from EBS
station 366, Lake Macquarie, NSW, thereby dem-
onstrating its ecological stability. This would sug-
gest a high level of genetic integrity.
VARIABILITY.?Most of the specimens assigned
to this category are listed below, with notes on
anomalies and variables, mostly to show the dif-
ficulty in treating this material as a cohesive
representation of a species.
Male "a," 2.98 mm, 1 dactyl 5 with only 5
spines, other with 5 spines and setule paired to
second proximal spine, telson entire or irregularly
emarginate, but not U. pentinus or U. pulcus be-
cause uropod 3 of pulcus form and pereopod 7
conformative to 872 form. CPBS 24N/4.
Male kkb," 2.70 mm, each dactyl 5 with 1 seta
accessory to proximalmost spine; telsonic cleft
gaping; primary flagellum thick. CPBS 24N/1.
Male "d," 2.48 mm, only 1 dactyl 5 present,
bearing 1 setule, pereopod 7 conformative, tel-
sonic formula with supernumerary short seta.
CPBS 24N/1.
Male "m," 3.65 mm, left dactyl 5 with 4 spines
and 2 setules (terminal and secondmost proxi-
mal), primary flagellum slightly thickened, telson
poorly cleft but each lobe with thick, fully ter-
minal spine-seta, outer ramus of uropod 3 with
only lateral spines (not setae), peduncle of uropod
2 with 5 lateral spines = 1-1-1-1 + 1. EBS 366.
Male "p," 2.55 mm, dactyl 5 with 4 spines,
second most proximal with small accessory spi-
nule, telson gaping. CPBS 24N/872.
Male "q," 2.70 mm, dactyl 5 with 4 spines, 3
setules, primary flagellum thick, telson gaping.
CPBS 24N/872.
Female "c," unmeasured, dactyls of pereopod
5 missing but telson deeply cleft and pereopod 7
conformative. CPBS 24N/1.
Female "e," ovigerous, 2.80 mm, dactyls of
pereopod 5 missing, telson deeply cleft, pereopod
7 conformative. CPBS 24N/1.
Female "g," unmeasured, dactyl 5 with 4
spines, setule on second proximal and terminal
only, pereopod 7 conformative, telson deeply
cleft, gaping. CPBS 24N/872.
Young female "k," 2.05 mm, dactyl 5 with 3
spines, 1 accessory spinule between apical and
next proximal spine, telson cleft. CPBS 24N/872.
Female "1," 2.90 mm, each dactyl 5 with 4
spines, 2 proximal spines, each with accessory
setule, but 1 side with third setule; telson abnor-
mal, entire. CPBS 24N/872.
Young female "t," 2.45 mm, dactyl 5 with 4
spines: 2 setae on 1 side, 1 terminal seta on
other side; telson gaping (damaged). CPBS 26N/
2.
Female v'u," unmeasured, dactyl 5 damaged,
all but 1 spine lost, 3 setules present, telson cleft,
pereopod 7 conformative, CPBS 24S/3.
Stations 967, 973, 975 (11 samples, received for
examination after above analysis). Most of the 44
NUMBER 360 61
individuals of both sexes conform to the general
pattern of morph 872, having gaped telsonic lobes
with apical setae and 1 or more setae in the
dactylar armament of pereopod 5. In a few spec-
imens the "setae" are thickened and spinelike,
though thinner and shorter than the spine they
accompany. In female "r," from sample 975/2,
both setae and additional spines are present.
PPBES.
VOUCHER MATERIAL? CPBS 24N/1: female
"h," 2.55 mm; CPBS 24N/4: male "a," 2.98
mm (illus.); CPBS 24N/872: female "1," 2.90
mm (entire telson); male "n," 2.70 mm (illus.);
PPBES 975/2: female "r"; EBS 366: male
"m," 3.65 mm. Others cited under "variability"
are also labelled and segregated but are subsidi-
ary.
MATERIAL.?CPBS, 5 samples from 3 stations
(15); PPBES, 11 samples from 3 stations (44);
EBS, 1 sample from 1 station (2).
DISTRIBUTION.?Victoria, Western Port to Port
Macquarie, NSW, 2-13 m, sand, sand and shell,
sand and some mud.
Urohaustorius pentinus, new species
FIGURES 23 (part), 24
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 mixing spines and setae. Armament
row on article 4 of antenna 2 even. Accessory
flagellum reduced. Main flagellar articles of male
antenna 1 thickened, aesthetascs 2-many per ar-
ticle, sometimes present on basal articles, article
5 of young male not bulbous. Coxa 1 not larger
than coxa 2, both bearing only tiny setules. Spines
on dactyl of pereopod 5 set singly in tandem.
Dactyls of pereopods 6-7 well developed. Article
2 of pereopod 7 with 2-3 stout posteroproximal
spines and scattered setules. Male sternite 6 with
apically blunt posteriorly directed ventral process.
Epimeron 2 bearing small tooth and 3-4 facial
setae in horizontal row. Tooth of epimeron 3
obsolescent. Inner rami of uropods 1-2 much
shorter than outer rami. Dorsolateral margin of
peduncle on uropod 2 with stout spines. Apico-
lateral corner of peduncle on uropod 3 with stout
spines; article 1 of outer ramus broad, with lateral
spines and setae, article 2 short and broad. Telson
emarginate, occasionally almost truncate, api-
cally scalloped, setae attached between scallops.
DESCRIPTION OF FEMALE.?Holotype female
"w," 2.28 mm: Many characters very similar to
U. pulcus and therefore illustrations reduced.
Head about 70% as long as wide, rostrum about
14% as long as remainder of head. Primary fla-
gellum of antenna 1 with 7 articles, accessory
flagellum 5-articulate. Facial formula of mixed
spines and setae on article 1 of antenna 1 = E,
5S, 2P; on article 4 of antenna 2 = 21, divided
weakly into 2 groups, spines dominant; on article
5 = 14; flagellum with 7 articles. Article 5 of
antenna 2 about 43 percent as wide as long, 63%
as long as article 4. Right incisor simple, left with
2 teeth, choppers on molars with 3 cusps; formula
on right palp = 8-1-3-1. Inner plate of maxilla
1 with 1 short apical seta, 1 large apicomedial
seta tightly adjacent to apical seta; outer plate
with 11 spines. Inner plate of maxilla 2 with 4
setae in weakly submarginal row. Inner plate of
maxilliped with 7 medial setae, outer plate with
1 apicolateral seta; palp article 2 with 6 setae in
facial row, article 3 with 2 setae in facial row.
Coxa 1 broadly subtriangular, apex blunt; coxa
2 similar; coxa 3 with 5 long ventral setae, none
posteriorly; coxa 4 with 5 ventral and 5 posterior
setae. Gills like U. pulcus.
Gnathopods like those of U. pulcus but article
5 of gnathopod 2 stouter.
Setal and spine formulas on pereopod 3 = 4, 5,
2-1-1, 4 + 4 + 1; on pereopod 4 = 3, 4, 2-1-1,
4 + 5 + 1, serrations like U. pulcus. Article 2 of
pereopods 5-7 progressively with medium to
sparse setal densities posteriorly, article 2 of per-
eopod 5 lacking mediofacial brush, dactyl with 3
spines in tandem; article 2 of pereopod 7 with
dorsoposterior armament in form of thickened
spines; shape of article 2 on pereopod 7 distinctive
(Figure 24 PI).
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental formulas =
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
w
FIGURE 24.?Urohaustorius pentinus, new species, holotype, female "w," 2.28 mm (n = male "n,
2.54 mm; u ? male "u," 1.88 mm;^ = female "y," 2.08 mm).
NUMBER 360 63
15-10, 13-9, 13-11; basal setal formulas = 8-1-
1-3,6-1-1-2, 7-1-1-2.
Epirneron 2 with 4 setae in oblique facial row.
Peduncle of uropod 1 with 6 lateral marginal
setae and 2 apicolateral setules; of uropod 2 with
3 evenly distributed lateral spines and 1 apicolat-
eral setule; dorsal setae = 2 and 2; setae of rami
= 6-3 and 6-3; inner rami extending 50% and
50% along outer. Apicolateral corner of peduncle
on uropod 3 with 5 spines, medial margin naked,
apicomedial corner with 1 setule, ventromedial
apical row(s) with 5 mixed setae; spine-seta for-
mula on lateral margin of article 1 on outer ramus
= S-SE-SE.
Telson about 1.2 times as wide as long, cleft
obsolescent, setal formula = 2P, 1, P, 1.
Lateral gland clusters well developed in pleon-
ites 3-5, weak in pleonite 6 and peduncle of
uropod 3.
Female "y," 2.08 mm: Article 4 of antenna 2
with 19 facial elements (6 and 13); coxa 4 with 4
ventral and 2 posterior setae, coxa 5 with 3 pos-
terior setae (2 in holotype), coxa 6 with 2 ante-
roventral setae (1 in holotype), 1 posteroventral
seta (2 in holotype); epimeron 2 with 3 facial
setae; epimeron 3 with main posteroventral setule
slightly above corner (holotype with this setule
below corner); apicolateral corner of peduncle on
uropod 3 with 4 spines; telsonic formula = 2P, 1,
P, 2 (illus.).
Female "x," 2.06 mm: Coxa 3 with only 3
setae; dactyl of pereopod 5 with only 2 spines;
article 2 of pereopod 7 with 3 dorsoposterior
spines; telson with same setule formula as holo-
type but cleft like female "y."
DESCRIPTION OF MALE (male "n," 2.54 mm).?
Like female but with following variations, mostly
nonsexual: Primary flagellum of antenna 1 with
8 articles, these scarcely thickened and not prolif-
erate; accessory flagellum 5-articulate. Article 4
of antenna 2 with 8 and 11 elements, article 5
about half as wide as long, and 70% as long as
article 4, with 11 elements; flagellum with 7
articles. Each molarial chopper with 3 cusps;
right and left formulas of palp article 3 = 7-1-3-
1 and 7-1-2-1. Coxa 3 with only 5 setae, anterior
lobe of coxa 6 with 2 setules. Spine and setal
formulas of pereopod 3 = 4, 4, 2-1, 4 + 4 + 1; of
pereopod 4 = 3, 4, 2-1-1, 4 + 4 + 1. Dactyl of
pereopod 5 with 3 spines; article 4 of pereopod 6
with only 4 posterior setae; article 2 of pereopod
7 with 3 posterodorsal spines. Formula of seg-
ments on rami of pleopods 1-3 = 15-11, 13-9,
13-10 (inner proliferate), setal counts not distinc-
tive. Epimeron 2 with 4 facial setae; epimeron 3
as stated for female "x." Uropods 1-2 like female
but inner ramus of uropod 1 with 4 setae (illus.)
and peduncle of uropod 2 with only 1 lateral
spine and 1 dorsal seta. Apicolateral corner of
peduncle on uropod 3 with 4 spines and 1 setule.
Telson like holotype.
INTERSEXES.?Four specimens (2.0-2.63 mm)
from the type-locality have short penial processes
and partly grown brood plates without setae.
HOLOTYPE.?NMV, female "w," 2.28 mm (il-
lus.).
TYPE-LOCALITY.?CPBS 12S/4, Western Port,
Victoria, 18 Mar 1965, 2.8 m, fine sand, some
mud and weed.
VOUCHER MATERIAL.?Type-locality: male
"n," 2.54 mm (illus.); male "q," 2.27 mm; male
"u," 1.88 mm (illus.); female "x," 2.06 mm (il-
lus.); female "y," 2.08 mm (illus.); 2 intersexes,
2.0 mm and 2.63 mm.
RELATIONSHIP.?This species is much smaller
than U. pulcus and the telson is emarginate rather
than cleft, with shorter, finer armament. The
presence of 2-3 stout proximoposterior spines on
article 2 of pereopod 7 is in contrast to the setae
found in U. pulcus; the shape of article 2 is rather
different. Both articles of the outer ramus of
uropod 3 are, by comparison, relatively shorter
and broader than those of U. pulcus. Setae on the
inner plate of maxilla 1 are more closely contig-
uous.
MATERIAL.?CPBS, 30 specimens from I sam-
ple.
DISTRIBUTION.?Victoria, Western Port, 2.8 m,
fine sand, some mud, weed.
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOC5Y
Urohaustorius merkanius, new species
FIGURES 25 (part), 26 (part)
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 composed of thin
elements; on article 4 of antenna 2 mixing spines
and setae. Armament row on article 4 of antenna
2 even. Accessory flagellum not reduced. Main
flagellar articles of only terminal male antenna 1
thickened, usually aesthetascs 2 per article and
usually absent on articles 1-3, article 5 of male
not bulbous. Coxa 1 not larger than coxa 2, both
bearing only tiny setules. Spines on dactyl of
pereopod 5 set in pairs on at least 1 locus. Dactyls
of pereopods 6-7 well developed. Article 2 of
pereopod 7 with 2 long posteroproximal setae,
sometimes a third towards midmargin, and se-
tules. Male sternite 6 with narrow, acute, poste-
riorly directed ventral tooth. Epimeron 2 bearing
medium tooth, with about 3 facial setae in hori-
zontal row. Tooth of epimeron 3 medium. Inner
rami of uropods 1-2 much shorter than outer
rami. Dorsolateral margin of peduncle on uropod
2 with or without spines. Apicolateral corner of
peduncle on uropod 3 with stout spines, outer
ramus with setae, with or without spines laterally
on article 1. Telson entire, distally crenulate.
ADDITIONAL CHARACTERS.?Article 5 of an-
tenna 2 oblong in adult females, more clavate in
males, peduncle of uropod 1 more than half as
wide as long.
GENERAL DESCRIPTION.?In contrast to U.
parnggius, the majority of adult specimens of U.
merkanius with (1) more numerous spines (of me-
dium thickness and length) on article 1 of antenna
1 (generally 5-8 elements); (2) primary flagellum
of male antenna 1 with slender articles bearing
only 2 aesthetascs each, and these absent on
articles 1-3 (except on 1 male, P.23354, 1500 m
E of Belmont Beach, NSW, presumably termi-
nal); (3) article 4 of antenna 2 with facial arma-
ment evenly aligned; (4) dactyl of pereopod 5
broad, with spines doubled at 1 or more loci; (5)
posterior margin on article 2 of pereopod 7 with
fewer long setae, generally 2 (less commonly 3),
confined to proximal half; (6) peduncle of uropod
1 stout and stumpy, more than half as broad as
long; (7) ordinary shape of article 2 on outer
ramus of uropod 3; and (8) sternal tooth on male
pereonite 6 distally attenuate and acutely
pointed.
DESCRIPTION OF MALE.?Holotype male "o,"
3.49 mm (Western Port; probably not
terminal): Head about 70% as long as wide,
rostrum about 15% as long as remainder of head.
Primary flagellum of antenna 1 with 10 unthick-
ened articles, 2 aesthetascs each on articles 4-8, 1
on article 9; accessory flagellum 7-articulate; fa-
cial formula of mixed spines and setae on article
1 of antenna 1 = 5E, 2P; on article 4 of antenna
2 = 2P, P, SSESEES-PSSESESSESESSESS; on
article 5 = 4SESESEESEESESSE; spines domi-
nant only on antenna 2; article 3 with 5 facial
setae; article 5 about 68% as wide as long, 61% as
long as article 4; flagellum with 9 articles. Right
incisor simple, left with 2 teeth, choppers on
molars with 4-5 cusps; formula on right palp
article 3 = 8-1-3-1, left = 7-1-3-1. Inner plate
of maxilla 1 with 1 short apical seta, 1 large
apicomedial seta; outer plate with 11 spines (sev-
eral with 3 side teeth instead of 2 as shown for U.
pulcus). Inner plate of maxilla 2 with 6 setae in
medial row. Inner plate of maxilliped with about
6 medial setae, outer plate with 1 apicolateral
seta, palp article 2 with 6 setae in facial row,
article 3 with 2 setae in facial row.
Coxa 1 broadly subtriangular; coxa 2 similar
but apex blunter; coxa 3 with 9 ventral setae all
in posterior half, posterior margin with 2 setae
both ventral; coxa 4 with 7 long ventral setae and
10 mixed posterior setae.
Setal and spine formulas on pereopod 3 = 4, 5,
2-1-1-1-1, 5 + 5 + 1 (= aberrant case, normal
= 6 + 5 + 1); on pereopod 4 = 3, 5, 2-1-1-1-1,
6 + 5 + 1, posterior margin of articles 5-6 with
broken row of serrations. Article 2 of pereopods
5-7 progressively with medium to sparse setal
densities posteriorly, article 2 of pereopod 5 lack-
ing mediofacial brush, dactyl with 5 spines in
tandem plus 2 partners. Pereonal sternite 6 with
acute midventral projection posteriorly directed
NUMBER 65
Gls
FIGURE 25.? [ rohaustonus merkamus, new species, holotype. male "o," 3.49 mm. / '. parnggius, new
species (p = holotype, male "p," 3.32 mm). U. yurrus, new species (g = female "g," 2.20 mm).
U. species M (c = female "c," 1.90 mm).
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
bW
FIGURE 26. ? Urohaustorms merkamus, new species, holotype, female "o," 3.49 mm (a = female
"a," 3.47 mm; b = male "b," 3.29 m m ; / = female "f," 3.42 mm; h ? female "h," 3.26 mm). U.
parnggius, new species (z ? female "z," 3.79 mm). U. species M (c ? female "c," 1.90 mm). Y
= pleopod. Z = male sternal spike.
NUMBER 360 67
(illus.). Article 2 of pereopod 7 with 3 posterior
marginal setae.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental formulas =
17-12, 15-11, 15-13 (inner proliferate); basal
setal formulas = 11-2-4-5, 5-0-2-0, 7-2-1-3.
Epimeron 2 with 3 setae in horizontal row.
Peduncle of uropod 1 stumpy, more than half
as wide as long, with 6 setae, apicolateral corner
with 1 setule; on uropod 2 with 3 apicolateral
spines closely contiguous, plus 1 setule; dorsal
setae = 3 and 1; setae of rami = 7-5 and 7-4,
inner rami extending 65% and 42% along outer.
Apicolateral corner on peduncle of uropod 3 with
4 spines, medial margin with 2 weak setae in
tandem, ventromedial apical row(s) with 10
mixed setae (rows distinct, 7 and 3), spine/seta
formula on lateral margin of article 1 of outer
ramus = Ee-Ee-Ee; article 2 of outer ramus
about 80% as wide as long, about 28% as long as
article 1.
Telson about 0.95 as long as wide, entire, ex-
treme apex flat, basally glandular; setal formula
= 2P, 1, 1, P, 1, 1.
Lateral gland clusters well developed in pleon-
ites 3-6, more weakly developed in peduncle of
uropod 3.
Male "b," 3.29 mm (Towra Point, Botany Bay,
NSW): Armament of articles 4-5 of antenna 2
ordinary; formula of article 4 = PP-SE-PSSSE-
SEESS-PSEESEESESESESS. Dactyls of pereo-
pod 5 stout, right with 1, left with 2 spines
doubled. Posterior margin of article 2 on pereo-
pod 7 with 3 setae, confined proximally. Epime-
ron 2 with 3 facial setae on right, 4 on left (illus.).
Peduncle of uropod 1 more than half as broad as
long, with 6 lateral setae; inner ramus unusually
short (only 56% as long as outer). Peduncle of
uropod 2 with 3 apicolateral spines and 1 lateral
spine (illus.); dorsal peduncular setae = 3 and 2;
ramal setae = 7 and 4, 7 and 3. Lateral spine-seta
formula on outer ramus of uropod 3 = ES, ES,
Ee, Ee/ES, Ee, Ee, Ee. Telson (illus.) apically
peaked, basally glandular; formula = 2P, 1, 1, P,
1, 1/2P, 1,P, 1, 1.
Male "p," 4.05 mm (offshore, 1500 m E of
Belmont Beach, 16 km S of Hunter R, NSW):
Article 1 of antenna 1, formula = 5 + 2P. Article
4 of antenna 2 formula = PP-ESP-SSSEESESS-
PSEESESESESESS; of article 5 = SSSE-S-ESS-
EESEEESS. Dactyl of pereopod 5 stout, spine
formula = 2-2-1-1-1/1-2-2-1-1-1. Article 2 of
pereopod 7 with 2 proximal posterior marginal
setae. Inner ramus of uropod 1 on left side unus-
ually long, 77% as long as outer. Peduncle of
uropod 2 with 1 long dorsal seta, 3 lateral and 3
apicolateral strong spines. Left and right spine-
seta formula for outer ramus of uropod 3 = Ee,
Ee, Ee, Ee/ES, Ee, ES, Ee, Eee. Telson tending
to asymmetry at apex, basally glandular; formula
= 1, 2P, 1, 1, P, 1, 1/2P, 1, 1, P, 1, 1.
Male "q," unmeasured, ~4 + mm (offshore
1500 m E of Belmont Beach, NSW): The only
specimen of U. merkanius with thickened basal
articles and many aesthetascs on main flagellum
of antenna 1 and reduced accessory flagellum.
Article 1 of antenna 1 with 6 spines + 2 penicil-
lates; article 2 with 7 fine spines, all short except
2 distal; main flagellum of 12 articles, aesthetasc
formula = 4, 4, 5, 4, 5, 5, 5, 4, 3, 1, 0; accessory
flagellum of 7 very thin articles. Article 3 of
antenna 2 with 5 setae; article 4 with normal U.
merkanius arrangement of armament, facial for-
mula = PPEEP-SSEESESESS-P-SSESEESE-
SESESS; article 5 about 63% as wide as long,
64% as long as article 4; formula = SSESESSE-
SEESESS. Dactylar spine formula for pereopod
5 = 1, 2, 2, 2, 1, 1/1, 3, 3, 1, 1. Article 2 of
pereopod 7 with 2 posterior setae confined to
proximal 1/3 of margin. Epimera 2 and 3 each
with small cusp and excavate margin; epimeron
2 with 5 setae in irregular oblique-horizontal
alignment and 1 ventrally, smaller, below the
middle seta. Peduncular spine formulas for pleo-
pods 1-3 = 2 and 1, 2 and 0, 2 and 0; segmental
formulas = 18-14, 16-12, 15-14; basal setal
formulas = 8-1-2-3 , 5-1 (setule)-1 - 1, 6-
l+2(setules)-l-3. Peduncle of uropod 2 with 2
lateral marginal spines close together, and 3 api-
colateral in close tandem. Peduncle of uropod 3
with 4 spines + 1 setule on dorsolateral apex;
lateral marginal formula of outer ramus on left
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
side = eS, ES, ES, Ee, Ee; article 2 extremely
short, 16% as long as article 1, and 75% as wide
as long. Telson longer than wide, extreme apex
flat, basally glandular; telsonic formula = 2P, 1,
1, P, 1, 1/2P, 1, 1, P, 1, 1 (very short).
DESCRIPTION OF FEMALE.?Female "f," 3.42
mm: Like male but with following variables:
Primary flagellum of antenna 1 with 9 articles,
accessory flagellum with 6 articles, article 1 of
peduncle with 5 thin setae plus 2 penicillate setae
in armament row. Antenna 2 as shown for female
of U. pulcus, article 5 about 48% as wide as long,
62% as long as article 4, facial armament formula
= PP-SE-PSEES-EESS-PSSESESSESESESS;
formula on article 5 = SEESS-EESEESESS.
Spine formula on posterior margin of article 5 of
pereopod 3 = 2-1-1; on article 6 of pereopod 4
= 5-4-1; otherwise like holotype. Fifth dactylar
spine formula = 2, 2, 1, 1, 1/1, 2, 2, 1, 1. Brood
plates thin, well setose. Uropod 1 peduncle with
only 2 dorsal setae, inner ramus with only 3 setae;
rami of uropod 2 with 7 and 3 setae. Ventrome-
dial setae on peduncle of uropod 3 in rows of 11
and 1. Telson basally glandular, apical crenula-
tion barely convex; setal formula = 2P, 1, P, 1,
1/2P, 1, P, 1, 1.
Female "a," 3.47 mm (Lake Macquarie Salts
Bay, NSW): Antennal armament in general
conforming to that of Victorian specimens; article
4 of antenna 2 formula = P-EE-PSSEESESEES-
PSEESESESESESS. Right and left dactyls of
pereopod 5 stout, each with 2 spines doubled.
Posterior margin of article 2, pereopod 7 with 2
proximal setae. Peduncle of uropod 1 more than
half as wide as long; inner ramus on left side
unusually long (75% as long as outer). Peduncle
of uropod 2 without spines. Telson apically con-
vex, basally glandular; formula = 2P, 1, 1, 1,
1/2P, 1, 1, P, 1, 1 (illus.).
Female %'m," unmeasured (1500 m E of Bur-
wood Beach, NSW): With slightly depressed
telsonic apex and telsonic formula as in male "p."
Female "c," unmeasured, and juvenile " j , " 2.35
mm (Merimbula, NSW): Both with telsonic
apex depressed. Dactyl of pereopod 5 with no
spines doubled on either side (juvenile only).
Otherwise these conform to U. merkamus diagnosis.
REMARKS.?Though there are recognizable dif-
ferences between Victorian and NSW forms of U.
merkanius as discussed below, they do not, on
present evidence, appear to be of specific signifi-
cance because of their extreme variability. All
Victorian specimens come from 1 sample,
WPBES 1714 (100% sand), just off the southwest
tip of French Island. Only 1 adult of the 71
Victorian specimens lacks spine doubling on at
least 1 locus of the dactyl on pereopod 5; while,
of the 17 subadults in the sample, 10 lack dou-
bling, 6 have 1 spine doubled on each dactyl
(bilateral), and 1 has doubling on 1 dactyl only.
Spines are not doubled on juveniles. Article 2 of
pereopod 7 most commonly bears 2 long setae
proximally, though occasionally 3 are present, as
in the holotype. The telson, in both NSW and
Victorian forms, shows some variation in precise
configuration, width to length ratio, and in the
number of marginal setae, but the extreme apex
is not as a rule peaked, and the apical crenulation
is usually flat or very weakly convex, and occa-
sionally, particularly in subadults, even slightly
depressed. Though the number and length of
telsonic setae varies, the true apex, whether con-
vex, flattened or depressed, is usually flanked by
2 very short, thin setae, next to which, more
laterally on each half of the telson is at least 1
much longer, stouter spine-seta, as illustrated in
male holotype and female "f," both from Victo-
ria, and female "a" from Lake Macquarie, NSW.
The condition in male "b," from Botany Bay, is
uncommon in either collection because the 2
apical setae are as long as the 2 subapical, and
the apical crenulation itself is strongly convex (as
in U. parnggius).
The offshore NSW individuals (28 males, 14
females, and 3 juveniles) from 28 samples are,
generally, larger than those from Victoria, with
an accompanying increase in the number of
spines and setae on some plates and appendages,
though article 2 of pereopod 7 carries only 2
posterior marginal setae. On the dactyl of the
fifth pereopod there are sometimes not 2 but 3
NUMBER 360
spines at the same locus, and the total number on
any dactyl is variable, as in the Victorian collec-
tion. Although no adult Victorian specimen bears
lateral spines on the peduncle of uropod 2 or on
article 1 of the outer ramus of uropod 3, some of
the NSW individuals do carry such spines. All
NSW adults examined, of both sexes, bear at least
I lateral spine on the peduncle of uropod 2; and
in 18 of them, mostly males, the outer rami of
uropod 3 also carry some spines as well as setae,
approaching in this respect the condition char-
acteristic of U. parnggius. The spination of these
NSW individuals is, however, vagarious. Spina-
tion is so erratic in occurrence and irregular in
distribution and numbers that no set pattern can
be discerned. No ramus has a spine at each locus
(unlike the stable condition found in U. parnggius,
where only the apex bears setae alone). In only
II of 42 adults examined are spines present on
both right and left rami, and then not necessarily
in corresponding number or position.
Article 2 on the outer ramus of uropod 3 is
shorter, compared with article 1, in NSW speci-
mens, than in the smaller Victorian specimens,
though it is still of about the same length-breadth
proportions. It has been observed in Urohaustonus
species that article 2 of uropod 3 is relatively
longer in smaller specimens, such as juveniles and
young subadults, than in the normally larger
adults of the same species.
The single adult female from Tasmania bears
1 lateral peduncular spine on uropod 2, but re-
sembles the Victorian form in the absence of
spines on uropod 3.
NSW estuarine specimens from Merimbula,
Botany Bay, and Lake Macquarie resemble the
offshore form in spination.
The male sternal process appears to be more
attenuate in the NSW specimens, but this may
well be artifact due to different preservational
techniques.
The occasional absence of the seta accompa-
nying the coupling hooks on the third pleopod of
some of the NSW males shows no pattern in
relation to other characters, except that it seems
to occur only in large males (though not in all
large males) and the seta is not present on the
only male with modified antenna 1, which is not
among the largest of them. It may be lost at
maturity.
HOLOTYPE.?NMV, male "o," 3.49 mm.
TYPE-LOCALITY.?WPBES 1714, Western Port,
Victoria, 10 Oct 1974, 0.3 m, sand.
VOUCHER MATERIAL.?Type-locality: female
"f," 3.42 mm (illus.); male "g," 3.30 mm; female
"h," 3.26 mm (illus.); EBS 148: male "b," 3.29
mm (illus.); EBS 366: female "a," 3.47 mm;
HDWBS: male "p," 4.05 mm; male "q," unmea-
sured ~4.0 mm; female "m," unmeasured; Day
and Kudenov coll., Merimbula NSW, female "c,"
unmeasured, and juvenile " j , " 2.35 mm.
RELATIONSHIP.?This species is easily distin-
guished from U. pulcus and U. pentinus in the
doubling of spines on the dactyl of pereopod 5;
and, unlike all specimens of U. pulcus and most
specimens of U. pentinus, the uncleft telson affords
a good distinguishing mark. In occasional speci-
mens of U. pentinus the telson is so nearly entire
that other characters, such as the presence of
stout spines on antenna 1, the shape and arma-
ment of uropod 3, the posterior margin of article
2 of pereopod 7, and the dactylar spination of
pereopod 5, may be used to distinguish the spe-
cies.
Absence of uropodal spines in the Victorian
form of U. merkanius at first appeared to segregate
the species clearly from U. parnggius, but the
erratic and variable occurrence of spines in the
NSW specimens renders the character valueless
in this respect. It should also be borne in mind
that all the Victorian specimens in hand come
from a single sample, and that additional mate-
rial from other localities could alter the present
picture.
Critical characters differentiating both Victo-
rian and NSW forms of U. merkanius from U.
parnggius are: (1) the arrangement of armament
on article 4 of antenna 2; (2) the presence or
absence of additional spines at 1 or more loci on
the dactyl of pereopod 5; and (3) the distribution
of posterior marginal setae on article 2 of pereo-
pod 7. The consistent occurrence, in combination.
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
of the sharp dorsoventral disjunction between the
most proximal group of elements and the main
section of the armament row on article 4 of
antenna 2, the single tandem arrangement of
dactylar spines on pereopod 5 (despite the large
body size), and the wide, even spread of setae
over the whole posterior margin of article 2 on
pereopod 7 (characters that have not been ob-
served together or severally in U. merkamus), offer
good grounds for assigning U. parnggius to a sep-
arate species.
In the open sea of Victoria and increasingly
more prominent towards Queensland, morphs or
sibling species prevail (U. parnggius, U. yurrus, U.
urungari) in which dactylar spines of pereopods
are arranged singly, in tandem.
MATERIAL.?WPBES, 1 sample (71); T.
Walker (Tasmania), 1 sample (1); EBS, 2 samples
(8); Day and Kudenov coll., Merimbula NSW
(2); HRDWB, 28 samples (44).
DISTRIBUTION.?Victoria, Western Port, to
Lake Macquarie, NSW, embayment and off-
shore, 0.3-28 m, fine sand, coarse sand, mud and
gravel.
Urohaustorius wingaro, new species
FIGURE 27 (part)
DIAGNOSIS. ? Dorsolateral armament rows on
articles 1 and 2 of antenna 1 bearing only thin
elements; article 4 of antenna 2 mixing spines
and setae. Armament row on article 4 of antenna
2 with sharp dorsoventral disjunction between
most proximal group of elements and main sec-
tion. Accessory flagellum reduced. Main flagellar
articles of male antenna 1 thickened, anesthetascs
2-many per article and present but usually absent
on article 1, article 5 of male bulbous. Coxa 1 not
larger than coxa 2, both bearing only tiny setules.
Spines on dactyl of pereopod 5 set singly in
tandem. Dactyls of pereopods 6-7 well developed.
Article 2 of pereopod 7 with 3 long posterior
setae. Male sternite 6 with broad rounded projec-
tion. Epimeron 2 bearing small tooth, with 2-3
facial setae in horizontal row. Tooth of epimeron
3 small. Inner rami of uropods 1-2 much shorter
than outer rami. Dorsolateral margin of peduncle
on uropod 2 with stout spine(s). Apicolateral
corner of peduncle on uropod 3 with stout spines;
outer ramus with setae and spines laterally on
article 1. Telson weakly to moderately cleft, lobes
gaping, bearing only 1 truly apical seta on each
lobe. Telsonic formula = 2P, 1, P, I.
ADDITIONAL CHARACTER.?Lateral elements of
article 1 on outer ramus of uropod 3 longer than
in U. pulcus and U. pentinus.
GENERAL DESCRIPTION.?In contrast to U. pul-
cus, body size of U. wingaro at maturity much
smaller (about 1.75 mm compared to 3.00 mm)
and article 2 of outer ramus on uropod 3 squat
and stout; in contrast to both (/. pulcus and U.
pentinus, armament group on article 1 of antenna
1 lacking spines; in contrast to U. merkamus, pe-
duncle and outer ramus of uropod 3 with lateral
spines; and in contrast to U. perkeus, coxa 3 poorly
setose.
DESCRIPTION OF MALE.?Holotype male "a,"
1.93 mm: Head about 75% as long as wide,
rostrum about 12% as long as remainder of head.
Primary flagellum of antenna 1 with 7 unthick-
ened articles, aesthetasc formula (proximal to
distal) = 0-1-2-3-2-1-0, accessory flagellum 5-
articulate. Facial formula of mixed spines and
setae on article 1 of antenna 1 = E, S, E, 2P; on
article 4 of antenna 2 in 4 sections, displaced at
second, as follows: P, P, EEP \ EEE-SEES-SSE-
SESS; article 5 swollen, 66% as wide as long, 66%
as long as article 4, formula = SSESESESS;
spines dominant only on antenna 2; article 3 of
antenna 2 with 2 facial setae; flagellum with 6
articles. Right incisor simple (but actually with 6
microcusps in waves), left simple, choppers on
molars with 3 cusps; formula on right palp article
3 = 7-1 -3 -1 , left (broken). Inner plate of maxilla
1 with 1 apical seta, 1 medium apicomedial seta;
outer plate with 11 spines. Inner plate of maxilla
2 with 4 setae in medial row. Inner plate of
maxilliped with 3 medial setae, outer plate with
1 apicolateral seta; palp article 2 with 3 setae in
facial row, article 3 with 2 setae in facial row.
Coxa 1 broadly subtriangular; coxa 2 similar
NUMBER 360 71
FIGURE 27.?Various species of Urohaustorius: Y ? ventral view of male apparatus; P = profile
of same. [r. wingaro, new species, holotype, male "a," 1.93 mm (c = male "c" 1.70 mm). U.
pulcus morph 872 (J = female "f," 2.90 mm; n = male "n," 2.70 mm). U. urungan, new species,
holotype, male "i," 2.40 mm (h = male "h," 2.55 mm). U. yurrus, new species, holotype, male
"r," 3.00 mm (o = male "o," 2.28 mm).
but apex blunter; coxa 3 with 3 ventral setae all
in posterior half, posterior margin with 1 setule
near corner; coxa 4 with 1 long ventral seta, 2
short setae, 2 setules and 4 posterior setules.
Setal and spine formulas on pereopod 3 = 3, 4,
2-1-1, 5 + 4 + 1 ; pereopod 4 = 3, 4, 2-1-1-1-1,
5 + 4 + 1; posterior margins of articles 5-6 with
broken row of serrations. Article 2 of pereopods
5-7 progressively with medium to sparse setal
densities (= 7, 6, 3 long and 5, 3, 2 short setae on
each article); article 2 of pereopod 5 lacking
mediofacial brush, dactyl with 4 spines in tan-
dem. (Note right pereopod 7 article 2 with 6 long
widely spread setae and no short setae.)
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental formulas =
15-10, 12-8, 13-10 (inner not proliferate): basal
setal formulas = 7-2-1-3, 2-1-1-1, 6-0-2-3.
Epimeron 2 with 3 setae in horizontal row.
Lateral margin of peduncle on uropod 1 with
6 setae, apicolateral corner with 1 setule; of uro-
pod 2 with 2 apicolateral closely contiguous spines
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
but no setule, plus 1 lateral spine; dorsal setae =
2-3 and 1; setae of rami = 8-3 and 7-2; ventral
peduncular medial setae = 1-2-1 and 3-2 (distal
to proximal); inner rami extending 50% and 50%
along outer. Apicolateral corner of peduncle on
uropod 3 with 2 spines (plus or minus a seta),
medial margin with no setae, ventromedial apical
row with 6 mixed setae, spine formula on lateral
margin of article 1 on outer ramus = SE-Ee.
Telson about 1.1 times as wide as long, cleft
about 0.20 of its length, setal formula = 2P, 1, P,
1.
Lateral gland clusters well developed in pleon-
ites 3-6, strongly in peduncle of uropod 3 and
weakly in pleonites 1-2.
PPBES male "b," 2.05 mm: With a few more
antennal armament elements than those from
Western Port; article 5 of antenna 2 swollen;
article 4 of pereopod 4 with only 3 apicodistal
long setae; pleopodal spine formulas = 2 and 1,
2 and 0, 2 and 0; segmental formulas = 14-9, 12-
8, 13-10; basal setal formulas = 5-2-1-3, 3-0-0-
1, 5-0-1-2; telsonic formula = 2P, 1, P, 1.
PPBES male "x," 2.15 mm: The only male
specimen found with thickened main flagellar
articles (3 basal), and proliferation of aesthetascs
on antenna 1 (formula = 3, 3, 4, 5, 5, 5, 5, 1, 0),
accessory flagellum reduced; article 5 of antenna
2 not as swollen as in other males, 55% as wide as
long and 67% as long as article 4; facial formula
for article 4 = P, EP | EEE-PSEESESESESS, for
article 5 = SEES-ESSEES. Spine-seta formula
for pereopod 3 = 3, 4, 1 (2)-1-1 ? 1, 4 + 4 + 1; for
pereopod 4 = 3, 3, 2-1-1, 4 + 4 + 1 on 1 side, 3,
3, 2-1-1-1, 4 + 4 + 1 on the other. Dactyl of
pereopod 5 with 3 spines. Peduncular formulas
for pleopods 1-3 = 2 and 1, 2 and 0, 2 and 0.
Epimeron 2 with 3 setae. Peduncle of uropod 3
with 3 apicolateral spines. Telsonic formula = 2P,
1, P. 1.
Male "s," 1.75 mm: Telson weakly incised
(like illustration of holotype) and peduncle of
uropod 2 with 1 fully lateral spine besides apico-
lateral spine, uropod 3 stubby, but articles 1-2 of
antenna 1 without stout elements, short elements
thin, article 2 of pereopod 7 with 3-4 posterodor-
sal stiff, stout but elongate setae, about 2-3 times
as long as those illustrated for holotype of U.
pentinus.
Male"m/' 1.77 mm: Telson moderately cleft,
closely similar to that of U. pulcus but article 2 of
pereopod 7 with enlarged setal-spines (illus.); uro-
pod 3 stubby, uropod 2 with 2 lateral spines (one
apical, one fully lateral); epimeron 2 with 2 facial
setae; articles 1-2 of antenna 1 with no fully
thickened short spines. Note striatc callus on me-
dial surface of article 5 on antenna 2 (illus.),
possibly typical of males in Urohaustoriidae.
Males "m" and "s" conform to description of
holotype, antenna 1 flagellum of each with 6
articles, accessory flagellum with 4; no aesthetasc
on basal article. Article 4 of antenna 2 in both
with dorsally displaced spine-seta row, formula
the same for each = SES-SES-ESESS. Spine
formula for pleopods 1-3 = 2 and 1, 2 and 0, 2
and 1.
REMARKS.?In all 5 Western Port males exam-
ined, basal articles of antenna 1 are not noticeably
thickened and article 5 of antenna 2 is stout and
swollen, not less than 66% as wide as long. Prob-
ably none is a mature male.
DESCRIPTION OF FEMALE.?Female "p," 1.62
mm: Formula on article 1 of antenna 1 = E, E,
2P; all elements on article 2 thin, 4 of these short
'and not plumose; primary flagellum with 6 arti-
cles, accessory flagellum with 4; aesthetasc for-
mula = 0-1-2-1-1-0. Article 3 of antenna 2 with
1 facial spine-seta, formula on article 4 = SE f
EE-PSES-SESESS; article 5 half as wide as long,
70% as long as article 4, formula = SES-ESES;
flagellum with 6 articles.
Long setae on coxa 3 = 4 ventral; on coxa 4
= 2 ventral, 2 posterior. Element formulas on
pereopod 3 = 3, 5, 2-1-1, 4 + 4 + 1; on pereopod
4 = 3, 4, 2-1-1-1, 4 + 4 + 1 . Setae of article 2 on
pereopod 5 = 1 0 long and 2 setules; on pereopod
6 = 7 long and 3 setules; on pereopod 7 = 3 long
stout setae and 1 setule in upper half, 3 setules in
lower half.
Epimeron 2 with 2 horizontally aligned setae.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental formulas =
NUMBER 360 73
13-9+, 11-7, 7+ -9; basal setal formulas = 4 - 1 -
2_3, 1-1-1-1, 5-0-1-3.
Peduncle of uropod I with 5 lateral setae, 1
dorsal seta, 1 apicolateral setule, rami with 6 and
4 setae; peduncle of uropod 2 with 1 apicolateral
spine, no lateral spines, 1 dorsal seta, rami with
6 and 2 setae. Telsonic formula = 2P, 1, P, 1.
(Uropod 3 and telson later lost in dissection.)
PPBES female "g," 1.5 mm: With setose
brood plates. Facial formula for main spine-seta
row on article 4 of antenna 2 = E-ES f SES-
SESESS on 1 side, SS-PSES t SES-SESESS on
the other; formula for article 5 = SESESES.
Article 4 of pereopod 4 with only 3 long setae
apicodistally. Dactyl of pereopod 5 with 2 spines
on right, 3 on left. Pleopodal peduncular formulas
for pleopods 1-3 = 2 and 1, 2 and 0, 2 and 1;
segmental formulas = 13-5+, 11-7, 12-6+; basal
setal formulas = 4-1-1-2, 1-0-1-1, 5-0-1-2 (5-
0-2-3). Telsonic formula = 2P, 1, P, 1.
Adult specimens of both sexes from Port Phillip
Bay conform to the specific diagnosis. They are
slightly larger than those from Western Port
(slightly over 2 mm), and most of the individuals
examined have 3 dactylar spines on pereopod 5.
In none of the 10 males examined for the char-
acter could a seta be detected accompanying the
coupling hooks on the peduncle of the third pleo-
pod, although this seta is present in females. In
both sexes there are only 3 long apicodistal setae
on article 5 of pereopod 4, compared with 4 in
Western Port specimens.
HOLOTYPE.?NMV, male "a," 1.93 mm (illus.).
TYPE-LOCALITY.?CPBS 12S/1, Western Port,
Victoria, 18 Mar 1965, 2 m, muddy sand, weed.
VOUCHER MATERIAL.?CPBS 01N/3: male
"m," 1.77 mm; CPBS 01N/4: female "p," 1.62
mm; male "s," 1.75 mm; CPBS 24N/872: male
"c," 1.70 mm (illus.); PPBES 919/2: female "g,"
1.5 mm; PPBES 920/5: male "b," 2.05 mm; male
"x," 2.15 mm.
RELATIONSHIP.?Urohaustorius wingaro has char-
acters in common with both the U. pulcus-U.
pentinus group (defined as couplet 12 of "Key I,"
page 51) and with the U. merkamus-U. parnggius
group (page 52, couplet 5). The cleft telson and
the presence of spines on the peduncle and outer
ramus of uropod 3 associate it particularly with
U. pulcus, and especially with morph 872, whereas
the absence of stout spines in the armament of
antenna 1 is characteristic of U. merkanius. The
cleft telson and the fact that it is very small at
maturity might suggest that U. wingaro is a neo-
tenic form of U. pulcus: and the absence of stout
spines on antenna 1 might support this view
because in small juveniles of U. pulcus they are
also absent. However, in such juveniles uropod 3
is more or less linguiform; article 1 of the outer
ramus is without lateral marginal spines, and is
not stout; article 2 is narrow and even longer,
relative to article 1, than it is in the adult. Uropod
3 of U. pulcus, therefore, is essentially different
from that of U. wingaro (with its short squat
spinose uropod 3).
U. wingaro is much closer in size to U. pentinus,
which it also resembles in the shape of uropod 3
and in the presence of spines on the outer ramus
and peduncle. But U. pentinus, even though of
small size, has well-developed stout spines in its
antennal armament; stout posteroproximal mar-
ginal spines on article 2 of pereopod 7 in place of
the long setae of U. wingaro; and its telson apically
scalloped and emarginate, not smooth and cleft.
The second antenna of nonterminal (?) males of
U. pulcus and U. pentinus is similar in shape to that
of the female, whereas in U. wingaro the male
article 5 is swollen. Adults of U. wingaro closely
resemble juveniles of U. perkeus in general appear-
ance, but may be distinguished by the larger
number and even spread of posterior marginal
setae on article 2 of pereopod 7 and the more
setose coxa 3 of that species.
U. wingaro and U. parnggius are the only 2
species described in the genus with distinctive
dorsoventral disjunction between the most prox-
imal and the main sections of the armament row
on article 4 of antenna 2. In both, article 5 of the
subadult male antenna 2 is swollen and shorter
than that of the female. U. parnggius, however, is
very much larger (3.00-3.79 mm) than the tiny
U. wingaro, its telson is entire; the posterior margin
of article 2 on pereopod 7 has setae spread over
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
the entire length; and the arrangement of molar-
ial cusps is rather different (see U. parnggius).
The consistent combination of characters of
antennae, pereopod 7, uropod 3, and telson make
U. wingaro relatively easy to identify and suggest
that it is a valid species.
MATERIAL.?CPBS, 4 samples from 3 stations
(6); PPBES, 21 samples from 11 stations (85).
DISTRIBUTION.?Victoria, Western Port and
Port Phillip Bay, 2-12 m, sand, muddy sand,
sandy silt, silty clay.
Urohaustorius parnggius, new species
FIGURES 25 (part), 26 (part)
DIAGNOSIS.?Dorsolateral armament rows on
article 1 of antenna 1 mixing long and short
elements of medium thickness; article 4 of an-
tenna 2 mixing spines and setae. Article 2 of
antenna I with fine spines only. Armament row
on article 4 of antenna 2 with sharp dorsoventral
disjunction between most proximal group of ele-
ments and main section. Accessory flagellum re-
duced in terminal male only. Main flagellar ar-
ticles of male antenna 1 thickened in terminal
male only, aesthetascs 2-6 per article, usually
absent on article 1 except in terminal male; article
5 of male antenna 2 bulbous. Coxa 1 not larger
than coxa 2, both bearing only tiny setules. Spines
on dactyl of pereopod 5 set singly in tandem.
Dactyls of pereopods 6-7 well developed. Article
2 of pereopod 7 with 4-7 medium to long, widely
spread posterior setae, most distal usually directed
dorsad. Male sternite 6 with short erect blunt
tooth. Epimeron 2 bearing medium tooth, with
3-4 facial setae in horizontal row. Tooth of epi-
meron 3 medium. Inner rami of uropods 1-2
much shorter than outer rami. Dorsolateral mar-
gin of peduncle on uropod 2 with stout spines.
Apicolateral corner of peduncle on uropod 3 with
stout spines, outer ramus with setae and spines
laterally on article 1. Telson entire, peaked at
apex (armament not diagnostic).
ADDITIONAL CHARACTER.?Peduncle of uropod
1 less than half as wide as long.
GENERAL DESCRIPTION.?In contrast to U. mer-
kanius, adult specimens have (1) fewer and nar-
rower spine-setae on article 1 of antenna 1, gen-
erally only 4, alternating longer and shorter; (2)
main armament row on article 4 of antenna 2
with sharp vertical displacement proximally; (3)
narrower dactyl of pereopod 5, and spines on this
article not doubled; (4) more and evenly spread
posterior marginal setae on article 2 of pereopod
7, instead of 2 or 3 confined proximally as in U.
merkanius; (5) narrower peduncle of uropod 1, at
least twice as long as wide; (6) squatter article 2
of uropod 3; (7) peaked telson (usually, but not
in holotype).
DESCRIPTION OF MALE.?Holotype male "p,"
3.32 mm (probably subterminal): Armament
on article 1 of antenna 1 (= 4E, 2P) consisting of
4 elements of moderate thickness and length,
longer alternating with shorter, + 2 penicillates
(illustrated); article 2 setose, with 6 fine spines, 1
apical spine elongate; flagellum of 9 unthickened
articles, aesthetasc formula = 0, 0, 1,2, 2, 2, 2, 1,
0; accessory flagellum of 6 articles, not reduced.
Basal group of facial spines and setae on article
4 of antenna 2 shifted ventrad (illus.); formula
= P, P, EE, P | S-PSESSEES-ESESSESS (dis-
junct at arrow); article 5 swollen, 68% as wide as
long, 63% as long as article 4, facial formula = 3S
+ 3S-EES-ESS-ESS.
Mouthparts of ordinary Urohaustorius form; 4
cusps on molarial chopper set on lateral protu-
berance very distally, reaching chopper surface;
left mandibular palp article 3 formula = 7-1-
3-1.
Outer plate of maxilla 1 with 11 spines; inner
plate narrow, with 1 short, truly apical seta, 1
long subapical. Inner plate of maxilla 2 with 6
setae in medial row; outer with 1 apicomedial
seta and 2 apicomedial rows of 7 and 9. Article 2
of maxillipedal palp with 6 setae in facial row.
Article 6 of gnathopod 2 only 66% as long as
article 5.
Coxa 3 with 5 long ventral and 2 posterior
setae; coxa 4 with 5 long setae.
Spine and setal formula of pereopod 3 = 4, 5,
2-1-1, 5 + 4 + 1 ; of pereopod 4 = 4, 5, 2-1-1-1-
NUMBER 360 75
1-1,6 + 5 + 1. Dactyl of pereopod 5 narrow,
with 6 spines in tandem (illus.). Article 2 of
pereopod 7 with 6 long posterior marginal setae
evenly spread. Sixth pereonal sternite with blunt
tooth.
Formulas of pleopods 1-3 = 2 and 1, 2 and 0,
2 and 1; segmental formulas = 17-13, 14-11, 14-
13; basal setal formulas = 9-1-2-3, 3-0-1-1, 7-
0-2-3.
Epimeron 2 with 5 (left) and 4 (right) setae.
(Illustration from male "q.")
Peduncle of uropod 1 subsequal to outer ramus,
with 8 lateral and 4 dorsal setae. Right peduncle
of uropod 2 with 3 lateral spines in tandem, a
long plume and 3 apicolateral spines, as well as
3 dorsal setae; left with 2 large lateral spines and
a tightly contiguous third, no plume, and 3 api-
colateral spines; setae on rami of uropods 1 and
2 = 8-5 and 8-3. Uropod 3 stout; peduncle with
4 apicolateral spines and one setule and 8-9 long
setae towards medial distoventral margin; outer
ramus lateral marginal formula = S, ES, ES, Ee,
article 2 very short (about 17% as long as article
1) set at an angle directed outwards. Peduncles of
uropods 1 and 2 scarcely, of uropod 3 heavily,
glandular.
Telson entire, about as tall as broad, less
sharply narrowed towards apex than usual in this
species; telsonic formula = 2P, 1, P, 1, 1/2P, 2, P,
1, 1.
Male "s," 3.00 mm: Generally resembling the
holotype, but presumed terminal male, with an-
tenna 1 modified. Main flagellum of antenna 1
with 11 articles, accessory flagellum reduced, with
6 thin articles; aesthetasc formula = 2, 2, 6, 5,
(6), 5, 6, 4, 4, 4, 1, 0; article 2 with 5 fine, short
to medium spines plus 2 elongate apical fine
spines. Article 5 of antenna 2 relatively longer
and narrower than in other (presumed subter-
minal) males, nearly 70% as long as article 4 and
52% as wide as long; article 4 about 80% as wide
as long, facial formula = P, P-EP | 3ES-PSSE-
SESS-ESESESS. Only 1 dactyl of pereopod 5
present, with 5 spines in tandem. Article 2 of
pereopod 7 with 7 evenly spread, posterior mar-
ginal setae, most distal longest, and dorsally di-
rected. Pereonite 6 with stout, blunt non-atten-
uate mid-sternal process. Peduncle of uropod 1
longer than outer ramus, less than half as wide as
long; lateral marginal formula for article 1 = SE,
SE, EE; article 2 very short, as broad as long,
about 15% as long as article 1. Telson as wide as
tall, apically narrow, formula = 1, 2P, 1, P, 1, 1,
1/2P, 1, P, 1, 1, 1.
Male "o," 3.50 mm: Resembling holotype.
Article 2 of antenna 1 with 4 fine spines and 1
long apical spine; flagellum of 9, accessory flagel-
lum of 7 articles. Article 4 of antenna 2 about
82% as wide as long, facial formula = P, P, 3EP
| 6E-PSSESESESESESS; article 5 slightly swol-
len, about 60% as wide as long and 73% as long
as article 4, flagellum of 9 articles, formula = 7S-
ESS-ESS-ESS. Telsonic formula = 2P, 1, P, 1,
1/2P, 1, P, 1, 1.
DESCRIPTION OF FEMALE.?Female "f," unmea-
sured: Resembling male. Article 1 of antenna 1
with additional spine (=5) + 2P; article 2 with 5
medium-short fine spines; main flagellum of 9,
accessory flagellum of 6, articles; aesthetasc for-
mula = 0, 0, 0, 2, 2, 2, 1, 1, 0. Facial formula for
article 4 of antenna 2 = PEE(?P) \ EEE, S-PEE,
ES-ESES-ESSESSESS; article 5 about 48% as
wide as long and 75% as long as article 4, formula
= 6S-3S-ESS-ESS; flagellum of 8 articles. Spine
and setal formulas of pereopod 3 = 4, 5, 2-1-1-1,
6 + 5 + 1; of pereopod 4 = 4, 5, 2-1-1-2, 6 + 5
+ 1. Only complete dactyl of pereopod 5 with 5
spines in tandem. Article 2 of pereopod 7 on each
side with 5 long posterior marginal setae, fully
spread. Epimeron 3 posteriorly with short, acute
cusp. Peduncle of uropod 2 with 3 lateral spines
close together in uneven tandem and 3 apicolat-
eral in close tandem. Inner ramus of uropod 3 as
long as article 1 of outer ramus; article 2 of outer
ramus only about 15% as long as article 1. Telson
taller than wide, apically peaked, entire, nonglan-
dular; formula = 2P, 1, 1, P, 1/2P, 1, P, 1. 1, 1.
Female "z," 3.79 mm: Fifth dactyls missing
from both right and left sides, but other charac-
ters conforming to specific diagnosis.
Article 1 of antenna 1 with 4 thin spines + 2
penicillates in armament row; article 2 with only
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
3 very fine spines, no very long apical; primary
flagellum of 9 articles, accessory of 7. Facial for-
mula on article 4 of antenna 2 = P, P-4E f 5E-
PSEESESSESESSESS, strong dorsoventral dis-
junction at arrow between most proximal and
main armament groups; article 3 with 3 setae;
article 5 normal; flagellum of 8 articles. Large
medial seta on inner plate of maxilla 1 not set
into notch. Spine and setal formulas on pereopod
3 = 4^  5? 2-1-1, 6 + 6 + 1; on pereopod 4 = 4, 5,
2-1-1-1,6 + 5 + 1. Setae numerous on coxae 3-
4. Article 2 of pereopod 7 with 5 setae evenly
spread, distalmost dorsally directed. Epimeron 2
with 4 setae. Peduncular formulas of pleopods 1-
3 = 2 and 1, 2 and 0, 2 and 1; segmental formulas
= 18-4, 15-10, 15-13; basal setal formulas = 6-
2-3-3, 2-1-1-1, 6-0-1-3. Urosomal glands as
shown for holotype of U. merkanius. Peduncle of
uropod 1 longer than outer ramus, less than half
as wide as long; outer ramus spinose, formula =
ES, ES, ES, Ee. Telson entire, strongly peaked
apically, formula = 2P, 1, P, 1, 1/2P, 1, P, 1, 1.
HOLOTYPE.?NMV, male "p," 3.32 mm (illus.).
TYPE-LOCALITY.?WPBES 1742/5, western en-
trance, Western Port, Victoria, 25 Nov 1974, 10
m, coarse-very coarse sand.
VOUCHER MATERIAL.?Type-locality: male
"u," 2.92 mm; male "q," 3.56 mm; WPBES
1743/1: female "f," unmeasured; CPBS, 24S/1:
female "z," 3.79 mm (illus.); HDWBS NSW, AM
P.23352: male "s," 3.00 mm; AM P.23362: male
"o," 3.50 mm.
RELATIONSHIP.?Urohaustonus parnggius appears
to be aligned with the U. merkanius group of
species by virtue of the uncleft telson and lack of
short, thick spines on articles 1 and 2 of antenna
1; and as in U. merkanius, article 5 of antenna 2 in
(subterminal?) males is broader and rounder than
that of the female. However, the single tandem
alignment of spines on the dactyl of pereopod 5,
the sharp disjunction of the proximal end of the
main armament row on article 4 of antenna 2,
and the full spread of posterior marginal setae on
article 2 of pereopod 7 segregate it from U. mer-
kanius; and the 2 latter characters from U. yurrus
and U. urungan. The dactylar spination of pereo-
pod 5 is characteristic of the U. pulcus group, but
the entire telson, lack of thick spines on article 2
of antenna 1, and armament of article 4 of an-
tenna 2 clearly distinguish U. parnggius from U.
pulcus and U. pentinus. The sharp displacement of
the proximal portion of the main armament row
on article 4 of antenna 2 is a character found
elsewhere only in U. wingaro of all the Urohaustonus
species described. From this species, however, U.
parnggius differs not only grossly in size, but in its
entire telson (compared with the gaping cleft in
U. wingaro), and in the greater number and full
spread of the posterior marginal setae on article
2 of pereopod 7.
MATERIAL.?CPBS, 1 sample (1); WPBES, 2
samples from 2 stations (4); HDWBS, NSW, 2
samples (2).
DISTRIBUTION.?Victoria, Western Port to
southern NSW, 10-18 m, fine to coarse sand.
Urohaustonus yurrus, new species
FIGURES 25 (part), 27 (part)
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 mixing spines and setae. Armament
row on article 4 of antenna 2 even. Accessory
flagellum not reduced. Main flagellar articles of
male antenna 1 thickened, aesthetascs 2-5 per
article and present on basal articles, article 5 of
male not bulbous. Coxa 1 not larger than coxa 2,
both bearing only tiny setules. Spines on dactyl
of pereopod 5 set singly in tandem. Dactyls of
pereopods 6-7 well developed. Article 2 of per-
eopod 7 with 2 (rarely 3) posteroproximal setae.
Male sternite 6 with short, broad, blunt ventral
protuberance. Epimeron 2 bearing blunt, small
to obsolescent tooth, with 2-3 facial setae in
horizontal row. Tooth of epimeron 3 small to
obsolescent. Inner rami of uropods 1-2 much
shorter than outer rami. Dorsolateral margin of
peduncle on uropod 2 with stout spine(s). Api-
colateral corner of peduncle on uropod 3 with
stout spines, outer ramus with setae and spines
laterally on article 1. Telson entire, bearing short
apical setules.
NUMBER 360 77
GENERAL DESCRIPTION.?In contrast to U. urun-
gan, more and stouter spines on article 1 of an-
tenna 1, especially in female (generally E, 3S,
2P); more spines on article 4 of antenna 2, espe-
cially in female; much narrower article 5 of an-
tenna 2 in males, both adult and subadult; nar-
rower article 6, and longer and thinner dactyl, of
pereopod 5; different form of male process on
sternite 6; article 2 of pereopod 7 with most
commonly 2 (exceptionally 3) proximoposterior
marginal setae, in contrast to the usual 3 (occa-
sionally 2) in U. urungan; greater number of spines
(usually 4, even in subadults) on the posterodorsal
corner of uropod 3 peduncle (U. urungari = 2,
exceptionally, 3); presence of spines as well as
setae on outer ramus of uropod 3 in females as
well as males; more sharply angled bulge medi-
ally at base of outer ramus; less massively glan-
dular, shorter, and apically much more broadly
rounded telson, reaching less than halfway along
the outer ramus of uropod 3.
DESCRIPTION OF MALE.?Holotype male "r,"
3.00 mm: Probably subterminal; head about
70% as long as wide, rostrum about 23% as long
as remainder of head. Primary flagellum of an-
tenna 1 with 10 articles, aesthetasc formula = 0-
0-1-2-2-2-2-2-1-0; accessory flagellum 7-artic-
ulate, not shortened; all short elements on article
2 of antenna 1 very thin. Facial formula of mixed
spines and setae on article 1 of antenna 1 = E,
3S, 2P; on article 4 of antenna 2 = P, PSE-
PSESESESE-SEESEESESESESESESESS; on
article 5 = SESESEESESESS; article 5 of an-
tenna 2 about 72% as long as article 4, 40% as
wide as long; article 3 with 3 facial setae; flagel-
lum 9-articulate.
Right incisor simple, left with 2 weak teeth,
choppers on molars with 4 cusps; formula on
right and left third palpar articles = 1-3-6-1.
Inner plate of maxilla 1 with 1 short apical seta,
1 large apicomedial seta; outer plate with 11
spines. Inner plate of maxilla 2 with 3 medial
setae. Inner plate of maxilliped with 4 medial
setae, outer plate with 1 apicolateral seta; palp
article 2 with 5 setae in facial row, article 3 with
2 setae in facial row.
Coxa 1 broadly subtriangular; coxa 2 similar,
but apex blunter; coxa 3 with long ventral setae
all in posterior half, posterior margin naked; coxa
4 with 2 long ventral setae, 3 setules and 4 long
or medium posterior setae and 5 setules.
Setal and spine formulas for pereopod 3 = 3, 5,
2-1-1-1, 5 + 4 + 1; on pereopod 4 = 3, 4, 2 - 1 -
1-1-1-1, 4 + 4 + 1; posterior margin of articles
5-6 with single row of mostly even large serra-
tions. Article 2 of pereopods 5-7 with progres-
sively medium to sparse setal densities posteriorly,
long setae = 8-4-2 respectively; article 2 of per-
eopod 5 lacking mediofacial brush, dactyl with 5
spines in tandem.
Peduncular spine formulas on pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental formulas =
17-11, 14-10, 14-12 (not proliferate); basal setal
formulas = 8-1-2-3, 2-1-1-1, 5-1-1-2.
Epimeron 2 with 3 setae in left horizontal row,
2 in right.
Lateral margin of peduncle on uropod 1 with
7 setae, apicolateral corner with 1 setule; of uro-
pod 2 with 1 disjunct lateral spine and 2 apico-
lateral spines; dorsal setae = 1 and 2; setae of
rami = 7-5 and 8-3; inner rami extending 58%
and 50% along outer. Apicolateral corner of pe-
duncle on uropod 3 with 3 spines (1 fewer than
normal for this species), medial margin with 1
weak seta; ventromedial apical row with about 8
mixed setae; spine formula on lateral margin of
article 1 of outer ramus = e, SE, SE, Ee; article
2 about 23% as long as article 1.
Telson about 1.1 times as wide as long, entire,
broadly rounded apically; telsonic formula = 1,
2P, 1,P, 1, 1.
Lateral gland clusters well developed in pleon-
ites 3-6, uropod 3 and telson.
Male "p," 2.38 mm: Presumably terminal
adult. Modified first antenna of 10 articles, basal
4 or 5 thickened, aesthetascs long, formula = 5,
4, 4, 5, 5, 5, 4, 2, 2, 0; reduced accessory- flagellum
of 6 thin articles. Article 1 of antenna 1 with 1
thin spine (spine-seta?) and 2 short, thicker spines
+ 2P; fine dorsal spines on article 2 = 6 short to
medium + 2 long apical. Article 5 of antenna 2
half as wide as long, 70% as long as article 4;
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
facial spine-seta formula for article 4 = P, P, E,
P-SESESSS-PSESESSESESESS/P, SESESSS-
PSSESESSESESS; for article 5 = SSE, SSE, SE,
SE, SS/SESESESESESS. Most proximal seta of
maxillary palp distinctly disjunct from 2 apical.
Setal and spine formula for pereopod 3 = 3, 5, 2-
1-1-1, 4 + 4 + 1; for pereopod 4 = 3, 5, 2-1-1-
2, 4 + 4 + 1.
Dactyl of pereopod 5 with 4/5 spines in tan-
dem, most proximal disjunct. Sternal projection
well developed, short, broad-based, blunt. Per-
eopod 7 with 2 proximal setae on posterior mar-
gin. Peduncular spine formulas of pleopods 1-3
= 2 and 1, 2 and 0, 1 (!) and 0; segmental
formulas = 14-10 (proliferating), l l(+)-9, 12-
10; basal setal formulas = 3(+)-2-l-3, 4-1-0-1,
5-1-1-3. Epimeron 2 with blunted cusp, 3 large
facial setae in horizontal row plus 2 much shorter
setae, 1 dorsal, 1 ventral, to the third seta. Epi-
meron 3 with well-developed small cusp, 3 ventral
marginal setae, posteriorly excavate, posterior
margin with 3 setules.
Peduncle of uropod 1 with 6 lateral, 6 medial,
and 3 large dorsal setae; outer ramus with 2 clear
apical and 1 subapical setae, in addition to mar-
ginals. Peduncle of uropod 2 with 2 midlateral
spines closely juxtaposed and 2 apical spines in
close tandem; outer ramus, in addition to mar-
ginals, with 1 clear central apical and 2 subapical
setae, I on each side. Peduncle of uropod 3 with
4 apicolateral spines; lateral spine-seta formula
on article 1 of outer ramus = ES, ES, Ee/ES, ES,
Ee; article 2 about 20% as long as article 1 and
80% as wide as long. Telson about as wide as
long, broadly rounded, glandular, extending
about 20%> along outer ramus of uropod 3; tel-
sonic formula = 2P, 1, P, 1, 1/2P, 1, P, 1.
Male "o," 2.28 mm (sternite illustrated): An-
tenna 1 flagellum of 10, accessory flagellum of 7,
articles; aesthetasc formula = 0-0-2-2-2-2-2-1-
l_0/0-0- l -2-2-2-2- l - l -0 . Article 5 of antenna
2 about 40% as wide as long and 48% as long as
article 4; flagellum of 8 articles. Epimeron 2 with
blunt cusp and 2 horizontal setae. Epimeron 3
with strong cusp.
Subadult male "s," 2.05 mm: With short pen-
ial processes and small sternite 6 projection. An-
tenna 1 flagellum of 8 articles; accessory of 5;
aesthetasc formula = 0-2-2-2-2-1-1-0. Article
5 of antenna 2 about 45% as wide as long and
74%) as long as article 4. Article 2 on outer ramus
of uropod 3 about 34%> as long as article 1. Telson
84%, as tall as wide; formula = 2P, 1, P, 1.
DESCRIPTION OF FEMALE.?Ovigerous female
"n," 2.40 mm: Formulas on article 1 of antenna
1 = E, 3S, 2P/E, 2S, 2P; all elements on article
2 very thin, 3 of them short; flagellum of 7
articles; accessory of 5; aesthetascs very long but
no more than 2 per article. Article 3 of antenna
2 with 2 facial spines; formula for article 4 (even)
= P, P, E-PSSESESS-PSESESSESEESESS; on
article 5 = SSESESESESESS.
Coxa 3 with 6 long ventral setae, 1 posterior.
Coxa 4 with 4 long ventral setae, 4 posterior.
Formula for pereopod 3 = 3, 3, 2-1-1,4 + 4 + 1;
for pereopod 4 = 3,4, 2-1-1-1,4 + 4 + 1 . Dactyl
of pereopod 5 with 4 spines in tandem, article 2
with 7 long posterior setae; of pereopod 6 with 4;
of pereopod 7 with 2 posterodorsal marginal se-
tae, slightly thickened.
Epimeron 2 with 2 setae.
Uropod 1, lateral setae of peduncle = 6 + 2
setules, dorsal = 2; rami = 6 and 4, left side
damaged. Uropod 2 peduncle with 1 small mid-
lateral spine plus 2 others of normal size, includ-
ing apical, all 3 in tandem; dorsal setae = 2(1);
ramal setae = 6 and 3. Peduncle of uropod 3 with
4 apicolateral spines: inner ramus like male; lat-
eral spine-seta formula = ES, EE, Ee/S, ES, Ee,
spines very fine, not short and thick. Telson al-
most as long as broad, formula = 2P, 1, P, 1.
Ovigerous female "g," 2.20 mm: Closely re-
sembling the above, but with 1 short plumose
seta fewer on article 2 of antenna 1 (illus.); 7
marginal setae on outer ramus of uropod 1; 1
lateral and 1 apical spine and 2 apical setae on
right peduncle of uropod 2; 3 apical spines on left
peduncle of uropod 3, 4 on right; spine-seta for-
mula of outer ramus of uropod 3 = ?-ES-EE/S-
Ee-Ee.
Female "m," 2.90 mm. With brood young.
This specimen is much larger than other oviger-
NUMBER 360 79
ous females in the collection but conforms to the
specific requirements. Armament row on article
1 of antenna 1 consisting of 2 fine spines (?spine-
setae) and 2 thick short spines + 2P; 5 very fine
spines on article 2 confined to distal 1/3 of article,
2 apical spines very long; main flagellum of 8
articles; unreduced accessory flagellum of 7;
aesthetasc formula = 0-0-2-1-2-2-2-0. Formula
of even armament row on article 4 of antenna 2
= P, P, EE-SEESESES-PSEESE-3SESESESS;
article 5 formula = 3SESES-ESESES3ES. Dactyl
of pereopod 5 with 5 spines in tandem. Epimeron
2 with 2 setae. Peduncle of uropod 1 with 7 lateral
and 2 dorsal setae. Left peduncle of uropod 2
with 1 stout lateral spine and 2 apical spines, and
in addition, more proximally, 2 short, middorsal
spines, 1 of moderate thickness, 1 very fine; right
peduncle with 2 midlateral marginal spines in
tandem, and in addition 2 short, proximodorsal
spine-setae. Ramal setae on uropods 1-2 = 9-6
and 8-4. Peduncle of uropod 3 with 3 apicolateral
spines on left side; right side damaged, with 2;
article 1 of outer ramus with sharply angled,
basal, medial bulge, lateral setal formula = EE,
ES, Ee/EE, E, (E)e, spines very fine; article 2
about 17 percent as long as article 1. Telson
sparsely glandular, broadly and evenly rounded
apically, reaching about 20% along outer ramus;
telsonic formula = 1, 2P, 1, P, 1/1, 2P, 1, P, 1.
BROOD JUVENILES (about 0.75 mm).?Notes
based on both juveniles in brood pouch of female
"m": Primary flagellum of antenna 1 with 4
articles, aesthetascs elongate; accessory flagellum
3-articulate; peduncular setae sparse. Article 3 of
antenna 2 with 1 long seta and 1 setule; article 4
with 2 long spines and 1 penicillate setule on
apical corner, ventral apex with 1 long plume, no
facial armaments; article 5 with 1 long and 1
penicillate setule at ventral apex, dorsal apex
with 1 long spine and 1 thin seta, no facials.
Articles 4 and 5 of pereopods 3 and 4 lacking
facial setae, article 4 with 1-2 apicoposterior se-
tae, article 5 with 1 pair of apicoposterior spines
only; formula on article 6 = 1 + 1 + 0 ; dactyl
much more elongate than in adult, spines highly
serrate. Dactyls of pereopods 3-7 very elongate
and with large sharp subapical thorn; no spines
on dactyl of pereopod 5. Articles 4-6 of pereopod
5 with apical sets of spines, but none set facially.
Inner rami of uropods 1-2 shorter than in adult,
each with only 1 apical seta, peduncle of each
with only 1 apicomedial seta total.
HOLOTYPE.?NMV, male "r," 3.00 mm (illus.).
TYPE-LOCALITY.?QM W.6386, station 58,
Middle Banks, Moreton Bay, Queensland, coll. S.
Cook, Sep 1972, 15.5 m, mud.
VOUCHER MATERIAL.?Type-locality: male
"p," 2.38 mm; female "n," 2.40 mm; subadult
male "s," 2.05 mm; QM W.6390: male "o," 2.28
mm (illus.); QM W.6384: female "m," 2.90 mm;
brood juveniles, ~0.75 mm; female "g," 2.20 mm
(illus.).
RELATIONSHIP.?Distinctions between U. yurrus
and U. urungan have already been enumerated in
the "General Description." These 2 species are
very close. They occur sympatrically in some
situations. Some of their distinctive characters
must be regarded as somewhat subjective (e.g.,
shapes of articles), and there are overlapping
characters such as setation of article 2 of pereopod
7, where U. yurrus has most commonly 2 (but
exceptionally 3), and U. urungari 3 (exceptionally
2) proximoposterior setae. The most important
distinguishing characters are: shape of the pro-
tuberance on sternite 6 of the male; shape of
article 5 of male antenna 2; and shape of the
telson.
In the collections there is no male of U. urungari
in which the flagellum of antenna 1 is modified
by thickening of basal articles and proliferation
of aesthetascs and so we cannot compare the 2
species in that terminal stage of development. All
males of the 2 species in hand show a difference
in shape of article 5 on antenna 2.
In all males of U. yurrus, whether with antenna
2 modified or not, article 5 is cylindrical to ovoid
in shape and about half as wide as long; in all 29
males of U. urungari this article is bulbous and
more than half as wide as long. No juvenile or
subadult male of U. yurrus examined shows a
structure similar to that of U. urungari on the sixth
sternite. Juvenile males of U. yurrus have a poorly
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
developed, wide-based, bluntly conoid process
with no indication of the low, bilobed protuber-
ance characteristic of U. urungan. Segregation on
the basis of telson shape alone shows that (1)
those individuals with broad, evenly rounded
telsons also have only 2 setae on article 2 of
pereopod 7; (2) article 6 of pereopod 5 is narrower
and the dactyl relatively longer than that of U.
urungan; (3) the peduncle of uropod 3 has 4
apicolateral spines; and (4) the armament of
antenna 2 is predominantly spinous (particularly
noticeable in the female). The alignment of the
armament row on article 4 of antenna 2 and the
setation of article 2 of pereopod 7 differentiates
both species from U. parnggius; and the entire
telson separates them from the clefted telson spe-
cies such as U. pulcus, U. wingaro, and U. perkeus.
The sporadic occurrence in both species of dou-
bling or pairing of dactylar spines on pereopod 5
is a further indication of a close relationship to U.
merkamus already obvious.
U. yurrus so closely resembles U. merkamus that
it might be regarded as a neotenic estuarine
morph of that species. The usual simple tandem
arrangement of the dactylar spines on pereopod
5 of U. yurrus is no bar to such a theory because
in small subadults and juveniles of U. merkamus
no doubling of these spines has been observed,
and some doubling in adults of U. yurrus and U.
urungan has been noted below (page 82). Apart
from this character, consistent distinctions be-
tween U. yurrus and U. merkamus are to be found
in
1. The shape and setation of the telson. In U.
yurrus the telson is broadly arched, and reaches
less than halfway along the outer ramus of uropod
3; in U. merkamus the apex is much narrower.
Adults and subadults of U.yurrus have only simple
seta between groups of penicillate setules. They
lack the second apicolateral pair of long stout
spine-setae, one on each side of the midpair. In
contrast, U. merkanius has 2 simple setae there. In
female "x," the apicolateral seta on each side is
already well developed, although the brood plates
are only about 1/3 grown and the 2 apical setae
are small.
2. The shape of article 5 on antenna 2 in the
male. This is cylindrical to ovoid in both adults
and subadults of U. yurrus, and less than half as
wide as long, but bulbous in all males of U.
merkanius examined except for the single adult
with modified antenna 1, in which it is narrower.
3. Shape of the male process on sternite 6;
short, broad, and bluntly conoid in both adults
and subadults of U. yurrus; fairly long, narrow,
hooked, and apically acute in U. merkanius.
MATERIAL.?Queensland, Moreton Bay, Mid-
dle Banks, 104 specimens from 5 stations.
DISTRIBUTION.?Queensland, Moreton Bay,
Middle Banks, 6-15 m, sand, mud.
Urohaustorius urungari, new species
FIGURE 27 (part)
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 mixing thin spines and setae. Arma-
ment row on article 4 of antenna 2 even. Acces-
sory flagellum not reduced. Main flagellar articles
of male antenna 1 not thickened, aesthetascs 2
per article and sometimes present on basal arti-
cles, article 5 of male bulbous. Coxa 1 broader
but not longer than coxa 2, both bearing only
tiny setules. Spines on dactyl of pereopod 5 usu-
ally set singly in tandem. Dactyls of pereopods 6-
7 well developed. Article 2 of pereopod 7 with 3
long stout posteroproximal setae and several se-
tules. Male sternite 6 with low transverse bilat-
erally lobed, bulbous ridge or flap. Epimeron 2
bearing small to obsolescent tooth, with 2-4 facial
setae in jagged horizontal row. Tooth cusp of
epimeron 3 obsolescent to small. Inner rami of
uropods 1-2 much shorter than outer rami. Dor-
solateral margin of peduncle on uropod 2 with
stout spine(s). Apicolateral corner of peduncle on
uropod 3 with stout spines; outer ramus with
setae and spines laterally on article 1. Telson
entire, apex extended, apical setules short.
GENERAL DESCRIPTION.?In contrast to U. yur-
rus, fewer and finer spines on article 1 of antenna
1, especially in female (generally 2E + 2P in
NUMBER 360 81
female, 3E 4- 2P in male); fewer spines and more
setae on article 4 of antenna 2, especially in
female; bulbous, not cylindrical, article 5 of an-
tenna 2 in the (nonterminal?) male; broader ar-
ticle 6 and relatively shorter, stouter dactyl of
pereopod 5; 3 (exceptionally 2) posterodorsal se-
tae on article 2 of pereopod 7 (U. yurrus, 2, excep-
tionally 3); only 2 (rarely 3) apicolateral spines
on peduncle of uropod 3 (U.yurrus 4); basal bulge
on outer ramus of uropod 3 less angularly defined;
telson taller and apically narrower, reaching far-
ther (at least halfway) along the outer ramus of
uropod 3.
DESCRIPTION OF MALE.?Holotype male "i,"
2.40 mm: Head about 65% as long as wide,
rostrum about 11% as long as remainder of head.
Primary flagellum of antenna 1 with 8 articles,
formula of aesthetascs = 0-0-2-2-2-1-1-0; ac-
cessory flagellum 5-articulate. Facial formula of
mixed spines and setae on article 1 of antenna 1
= E, S, 2P; on article 4 of antenna 2 = P, P, E,
P-ESEES-PSSEEESESESESS; on article 5 =
SESESESSESS; article 3 with 3 facial setae (2
setae and setule); flagellum with 6 articles.
Right incisor simple, left with 2 extremely weak
teeth; choppers on molars with 3-4 cusps; for-
mula on right palp article 3 = 6-1-3-1, on left
(= missing). Inner plate of maxilla 1 with 1 short
apical seta, 1 large apicomedial seta; outer plate
with 11 spines. Inner plate of maxilliped with 3
medial setae, outer plate with 1 apicolateral seta;
palp article 2 with 4 setae in facial row, article 3
with 2 setae in facial row.
Coxa 1 broadly subtriangular; coxa 2 similar
but apex blunter; coxa 3 with 4 ventral setae, 11
in posterior half, posterior margin with 1 seta
ventrad; coxa 4 with 4 ventral and 1 posterior
setae.
Setal and spine formulas on pereopod 3 = 3, 5,
2-1-1-1, 5 + 4 4- 1; on pereopod 4 = 4, 4, 2 -1 -
1-1-1-1,5 + 4 + 1; posterior margin of articles
5-6 with single row of mostly medium, even
serrations. Article 2 of pereopods 5-7 progres-
sively with medium to sparse setal densities pos-
teriorly (8-8-3), article 2 of pereopod 5 lacking
mediofacial brush, right dactyl with 4 spines in
tandem, left with 5 spines, 2 of these forming
pair!
Peduncular spine formulas of pleopods 2-3
(pleopod 1 missing) = 2 and 0, 2 and 1; segmental
formulas = 12-8 and 13-12 (inner proliferate on
pleopod 3); basal setal formulas = 4-0-1-1 and
7-1-2-3 (see specimen " j " below for complete
pleopodal formula).
Epimeron 2 with 4 setae in horizontal facial
row.
Peduncle of uropod 1, lateral margin with 6
setae, apicolateral corner with 1 setule; of uropod
2 with 1-2 disjunct lateral spines and 1-2 apico-
lateral spines plus setule, dorsal setae = 2 and 2,
setae of rami = 7-4 and 7-4, inner rami extending
60% and 60% along outer. Peduncle of uropod 3
short and stubby, more than half as wide as long,
subequal to, or shorter than, outer ramus, apico-
lateral corner with 2-3 spines, medial margin
naked, ventromedial apical row with 9 mixed
setae, article 2 of outer ramus about 33% as long
as article 1, about 80% as wide as long, armament
formula on lateral margin of article 1 = S-E(S?)-
Ee/ES-Ee.
Telson about 1.0 times as wide as long, entire,
setal formula = 2P, 1, P, 1 (midapical).
Lateral gland clusters well developed in pleon-
ites 3-6, more weakly developed in peduncle of
uropod 3, weak to moderately developed in tel-
son.
Male " j , " 275 mm: With well-developed ster-
nite 6 process (illus.). Facial formula on article 1
of antenna 1 = 3E + 2P. Spine-seta formula for
article 4 of antenna 2 = P, P-EE-P-ESEES-
PSEEEESESESESS; for article 5 = SESESE-
SESSS; flagellum of 6 articles. Dactyl of pereopod
5 with 5 spines in tandem, left side slightly ab-
normal, with spines 2 and 3 short. Article 2 of
pereopod 7 with 3 dorsoposterior setae. Epimeron
2 with 3 facial setae ( 2 + 1 socket); epimeron 3
with tiny cusp and slightly excavate posterior
margin. Peduncular spine-seta formulas for pleo-
pods 1-3 = 2 and 1, 2 and 0, 2 and 1; segmental
formulas = 15-10, 12-9(8), 13-12 (inner prolif-
erate); basal setal formulas = 8-1-2-3, 3-1-1-1,
7-1-1-3. Peduncle of uropod 3 with 3 apicolat-
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
eral spines, lateral marginal formula for outer
rami = ES, Ee/ES, Ee. Telson barely taller than
wide, apex flattened, setal formula = 2P, 1, P, 1/
2P, 1, P, 1.
Male "h," 2.55 mm: Article 1 of antenna 1
formula = E (long) EE + 2P. Spine-seta formula
on article 4 of antenna 2 = P, P-EE-PSEES-
PEEESEEESESESS; on article 5 = SSES-ESE-
SESS; flagellum of 6 articles. Dactyls on right
and left fifth pereopods each with 5 spines, none
doubled, though in 3 groups, 2-1-2 on 1 side.
Article 2 of pereopod 7 with 3 dorsoposterior
setae. See illustrations for sternite 6. Peduncle of
uropod 3 with 3 spines; lateral marginal spine-
seta formula of article 1 = ES, Ee/ES, Ee. Telson
barely taller than wide, peaked; telsonic formula
= left 2P, 1, P, I (unpaired mid apex)/right 1,
2P, 1,P.
DESCRIPTION OF FEMALE.?Large, ovigerous fe-
male "g," unmeasured, with 2 eggs in brood
pouch: Setal formula on article 1 of antenna 1
= 2E + 2P; spines on article 2 all very fine, 1
short, 2 medium, 1 long, all distal; flagellum
ordinary, with 7 articles; accessory flagellum with
5 articles. Spine-seta row on article 4 of antenna
2 even, formula = P, P, EE, P-5E-PEE-ES3ESS
(extreme reduction of spines); article 5 tending to
ovate in shape, about 60% as wide as long, 60%
as long as article 4, formula = S3eS3ESEESS
(abnormally large number of elements due to
increased number of setae). Right and left dactyls
of pereopod 5 each with 6 spines, distal 2 "paired"
(see note on p. 80) on each side. Right pereopod
7 with 3 dorsoposterior setae on article 2, left
missing. Posteroventral cusps on epimera 2 and 3
obsolescent. Epimeron 2 with 4 setae in jagged
row. Peduncular spine/seta formulas for pleopods
1-3 = 2 and 1, 2 and 0, 2 and 1; segmental
formulas = 17-12, 14-9, 16-15 (inner ramus pro-
liferate on both sides); basal setal formulas = 8-
(1 + 1 setule)-l-3, 2-0-1-1,7-0-2-(2+ 1 setule).
Peduncle of uropod 1 subequal in length to outer
ramus, much more than half as wide as long, with
8 lateral setae, 1 apicolateral setule, and 2 large
facial setae, in addition to ordinary medial ar-
mament; peduncle of uropod 2 with large dorsal
seta, laterally with 1 midmarginal spine and 2
apicolateral spines plus 1 spinule. Outer rami of
first and second uropods with triad of setae at
apex and 6 medial setae; inner rami of both
uropods about 60% as long as outer, each with 5
medial setae. Peduncle of uropod 3 glandular,
with 2 apicolateral spines; inner ramus extending
beyond apex on article 1 of outer ramus; formula
for lateral margin of article 1 = Ee, Ee/Ee, E;
article 2 about 30% as long as article 1. Telson
slightly taller than wide, heavily glandular, api-
cally narrow, setation symmetrical; formula =
2P, 1, P, 1/2P, 1, P, 1.
Female "n," 2.60 mm: Article 1 of antenna 1
with formula = E (long) + 2E + 2P; article 2
with 4 very fine spines, 1 short, 2 medium, 1 long,
all distal; flagellum of 7 articles, aesthctascs very
long, formula = 0-0-2-2-1-1-0; accessory flagel-
lum of 5 articles. Spine-seta formula on article 4
of antenna 2 = P-P-E-P-4ES-PSES3E SESESS;
article 5 cylindrical, though very slightly broader
towards distal apex, apex itself slightly rounded,
spine-seta formula = SSEESEESESS. Setal and
spine formulas on pereopod 3 = 5, 5, 2-1-1-1, 4
+ 4 + 1; on pereopod 4 = 4, 5, 2-1-1-1, 4 + 4
+ 1. Article 6 of pereopod 5 about 75% as wide as
long, only 1 dactyl present, with 5 spines in
tandem. Article 2 of pereopod 7 with 3 dorsopos-
terior setae. Cusps obsolescent on epimera 2 and
3; epimeron 2 with 2 and 3 setae. Peduncular
spine and seta formulas for pleopods 1-3 = 2 and
1, 2 and 0, 2 and 1; segmental formulas = 15-11,
13-9, 15-13 (inner proliferate); basal setal for-
mulas = 8-1-2-3, 3-0-0-1, 6-0-2-2. Peduncle of
uropod 1 bearing 6 (7) lateral, 2 large dorsal, and
5 medial setae, 1 medioapical medium-length seta
plus 1 setule; outer and inner rami each with 5
medial marginal setae. Peduncle of uropod 2 with
1 midlateral spine, 2 spines plus 1 spinule on
apex, and 1 large dorsal seta. Outer rami of first
and second uropods broad apically, with a triad
of setae. Peduncle of uropod 3 with 2 apicolateral
spines; article 1 of outer ramus on left side with
1 lateral marginal spinule, lateral margin on right
side naked; article 2 of outer ramus unusually
long, about 45% as long as article 1; inner ramus
NUMBER 360 83
extending beyond apex of article 1. Telson about
as wide as tall, glandular, asymmetrically setose;
telsonic formula = 2P, 1, P, 1, I (midapical)/l,
2P, 1, P.
Female "v," 2.75 mm: With symetrically se-
tose, tall, flat-apexed, sparsely glandular telson,
conforming to diagnosis and description of U.
urungan, but outer ramus of uropod 3 bearing
spines as well as setae (only known female so
equipped); formula for lateral margin of article
1 = ES, Ee/s, ES, Ee. Spine-seta formula for
article 4 of antenna 2 = P, P, P-EEEES-PSEEE-
SEESESESS. Left dactyl of pereopod 5 with 4
spines, right abnormal or damaged. Article 2 of
pereopod 7 with 3 dorsoposterior setae. Epimeron
2 with 2 setae. Outer rami of uropods 1 and 2
apically broad with 3 setae almost side by side.
Peduncle of uropod 3 with 2 apicolateral spines.
Female "f," 2.66 mm: Formula on article 1
of antenna 1 = E (long), E, 2P; all elements on
article 2 very thin. Article 3 of antenna 2 with 1
facial setal spine, formula on article 4 (even) = P,
P, P, P-ESEE-PEESEE-ESESESS, setae domi-
nant, on article 5 = SEESESESS, article 5 ovate
(not bulbous form of male).
Dactyl of pereopod 5 with 5 spines in tandem;
article 2 of pereopod 6 of thin form.
Peduncular spine formula of pleopod 1 = 2
and 1, segmental formula = 14-10, setal formula
of rami = 7-1-2-3 (to substitute for missing male
pleopod 1); other female pleopods badly dam-
aged.
Uropod 1 peduncle lateral setae = 6, dorsal =
2, rami = 8 and 4; uropod 2 peduncle lateral =
1 spine, apicolateral = 2 spines and setule, dorsal
setae = 2; ramal setae = 7 and 4. Uropod 3
peduncle apicolateral spines = 2, inner ramus like
male; setae formula on lateral article 1 of outer
ramus = e-E.
Telson as long as broad, armament formula =
2P, 1, P, 1 (unpaired middle).
REMARKS.?Many characters, such as numbers
and arrangement of spines and setae on append-
ages and exact configuration of structures such as
the telson, are variable in this species. Individual
abnormalities and malformations are not uncom-
mon (e.g., dactyl of pereopod 5). There are iso-
lated exceptions in almost all the characters gen-
erally used in the diagnosis of the species. How-
ever, characters of importance in the genus Uro-
haustorius, which are consistently stable in U. urun-
gari, are: absence of stout spines in the armament
of antenna 1; preponderance of setae over spines
on article 4 of antenna 2 in the female; bulbous
(sometimes almost spherical) shape of article 5 of
antenna 2 in the male (terminal male probably
unknown); absence of typical U. merkanius kind of
spine-doubling on the dactyl of pereopod 5;
unique form of the protuberance on sternite 6 in
males; absence of any long spine-seta on the
telsonic apex.
In all males in the collection article 5 of an-
tenna 2 is bulbous, and more markedly so in the
smaller individuals with an ill-defined sternite 6
process. In U. merkanius, U. parnggius, and U. win-
garo (all with bulbous article 5 in subterminal
males) this article is more slender in such males
as have been found with modified antenna 1,
which are assumed to be at the terminal stage.
This suggests that there may well be such a
terminal stage in U. urungari, though it is, as yet,
unknown.
Spines on the dactyl of pereopod 5 are rela-
tively longer and more slender than those of U.
merkanius or U. parnggius. The most proximal spine
is often shorter and decidedly disjunct; where
there are 5 spines present (the most usual condi-
tion in adults), the remaining distal 4 are fre-
quently in 2 groups. In cases where "pairing" or
"doubling" has been observed, as in some large
individuals and ovigerous females, it is, except in
1 recorded instance, the 2 most distal spines that
are affected. Instead of being close together in
tandem, they have assumed the "doubled" posi-
tion of being almost side by side horizontally.
This condition has involved simply a slight
change in position, and the paired spines are of
equal length, and not, as they are in U. merkanius,
in groups comprising 1 long spine plus 1 or more
shorter elements.
Females of U. urungari do not normally bear
spines laterally on article 1 of the outer ramus on
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
uropod 3 (female "v" is the only exception noted),
and frequently have no armament at all on this
article except at the apex. Even in ovigerous
females there may be only a simple spine-seta
laterally, not accompanied by a long plumose
seta. Most commonly there are only 2 apicolateral
spines, rather slender, on the peduncle of uropod
3. In a few large specimens of both sexes 3 spines
have been observed on 1 side (as on the holotype)
or both; but never 4 spines, as in U. yurrus.
Although some individuals have a bilaterally
symmetrical telson with a pair of setae, 1 on each
side of the apical midline, there is a strong tend-
ency in this collection of U. urungan towards loss
of 1 seta of the apical pair. In some specimens the
seta remaining retains its original position, cre-
ating apical asymmetry; in others it has become
truly median. Deviations in apical setation have
not so far been found to be associated with any
distinctive character, and occur in both sexes.
HOLOTYPE.?NMV, male "i," 2.40 mm (illus.).
TYPE-LOCALITY.?QM W.6386, station 58,
Middle Banks, Moreton Bay, Queensland, Sep
1972, 15 m, mud.
VOUCHER MATERIAL.?Type-locality: male "h,"
2.55 mm (illus. sternite, 2 figs.); female "n," 2.60
mm; female "g," unmeasured; male "k," 2.40
mm; male "1," 2.68 mm; QM W.6383: male " j , "
2.75 mm; QM VV.6385: female "f," 2.66 mm;
QM W.6392: female "v," 2.75 mm; QM W.6387:
female "o," 2.45 mm.
RELATIONSHIP.?The unique form of the male
process on sternite 6 of U. urungari is the most
important factor in its segregation from all other
species of Urohaustorius. Many of the other differ-
ences between U. urungari and U. yurrus, though
consistent, are subtle: telsonic configuration; ar-
mament on article 4 and shape of the male article
5 of antenna 2; shape of article 6 and the dactyl
of pereopod 5; setation of article 2 of pereopod 7.
Characters, such as setation and shape of the
telson and the exact spine/seta ratio on article 4
of antenna 2, are variable in both species (though
spines predominate on article 4 of antenna 2 in
U. yurrus and setae in U. urungari); and there is a
very small degree of overlap in some characters
(pages 79, 80). These criteria are difficult and
laborious to use in practical identification of eco-
logical survey material; and of course with limited
material we cannot assume that they are valid
specific differences. Nevertheless, they have been
replicated to our satisfaction, and there are only
a very few individuals in the collection that can-
not be confidently assigned, on the basis of these
criteria, to 1 group or the other.
MATERIAL.?Queensland, Moreton Bay, Mid-
dle Banks, 48 specimens from 6 stations.
DISTRIBUTION.?Queensland, Moreton Bay,
Middle Banks, 6-15 m, mud, sand.
Urohaustorius species M
FIGURES 25, 26 (part)
Known as the Malabar form, 4 specimens from
the Australian Shelf Benthic Survey off Malabar,
NSW, cannot be firmly identified with any of the
foregoing species in Urohaustorius. These speci-
mens, all females, appear, apart from the entire
telson, to have more in common with U. pulcus
and U. pentinus, and particularly the latter, than
with the U. merkanius group. The largest specimen,
female "x," 2.31 mm, has a broad telson narrow-
ing towards the apex, whereas the other 3 speci-
mens have the ordinary, mostly rounded, broad
telson; but in all 4 specimens the telson is apically
flattened. The presence of several short stout
spines on article 2 of antenna 1 appears to sepa-
rate these specimens from either U. merkanius or
U. parnggius, and the absence of doubled spine
positions on the dactyl of pereopod 5 and the
spinous article 1 on the outer ramus of uropod 3
remove the specimen from U. merkanius (unless
they are neotenous adults). The normal spination
pattern on article 4 of antenna 2 distinguishes the
specimens from U. parnggius and U. wingaro.
Distinctive characters of this morph are (1)
narrow article 2 of pereopod 6; (2) presence of 2
very long, stout, plumose setae at the proximal
end of the spine group on article 1 of antenna 1.
The inconsistency of the armament formulas
on article 1 of antenna I is, however, anomalous,
NUMBER 360 85
as seen in the following tabulation of female
specimens.
Specimen
"y
"d"
Antenna I
2E(long), 2S, 2P
2E, 2S, 2P
2E, S, 2P
2E, 2S, 2P
Opposite antenna 1
E(short), 3S, 2P
2E(long), 2E(short), 2P
E, 3S, 2P
2E, 2S, 2P
Only female "d" appears to be normal and
consistent with expectation in stability. One prox-
imal long seta has been observed in a specimen of
U. metungi (illus.); and 2, as in the Malabar form,
in 2 juveniles of what is probably U. pulcus from
Western Port, and in a sample of brood young of
U. yurrus from Queensland. In U. perkeus article 1
carries only such long setae. We consider the
possibility that this is a juvenile character and
that specimens of U. species M are mutant neo-
tenes. More specimens are awaited.
DESCRIPTION OF FEMALE.?Female "c," 1.90
mm: Armament row on article 1 of antenna 1
= 2E, 2e, 2P/2E, 2S, 2P. Armament row on
article 4 of antenna 2 with all elements aligned,
formula = P, PEE-PSESS-PSSESESESESESS.
Dactyl of pereopod 5 with 3 spines in tandem.
Epimeron 2 with 4 setae; tooth of epimeron 3
sharp. Uropod 1 as in typical specimens of U.
merkanius; peduncle with 5 lateral and 3 dorsal
setae; inner ramus not as short as usual. Peduncle
of uropod 2 with either 1 or 2 apicolateral spines
and 1 lateral spine; inner ramus shorter than in
U. merkanius; setae on rami of uropods 1 and 2 =
6 and 3, 6 and 2. Apicolateral corner on peduncle
of uropod 3 with 4 spines; article 1 of outer ramus
with only 2 sets of armaments. Telson truncate as
in specimen "u" of U. pentinus but not distinctly
excavate; setal formula = 2P, 1, P, 1.
Superfemale "x" of dwarf form, 2.31 mm:
Armament row on article 1 of antenna 1 with 2
long plumes, 4 short spines, 2 penicillate setules.
Armament row on article 4 of antenna 2 with
all elements aligned, formula = P-PEE-
PSESEESS-PSSEESEESESESESS; article 5 =
3SESESEESEES. Dactyl of pereopod 5 with 4
spines in tandem. Epimeron 2 with 4 setae. Pe-
duncle of uropod 2 with 3 apicolateral spines and
1 lateral spine. Telson entire, formula = 2P, 1, P,
1.
VOUCHER MATERIAL.?AM SBS E2S3 and
E2S4, female klx," 2.31 mm; female "c," 1.90 mm
(illus.); female "d," 1.75 mm; female "y," 1.73
mm.
MATERIAL.?SBS, 2 samples (4).
DISTRIBUTION.?NSW, off Malabar, ~20 fath-
oms (37 m), coarse sand.
Urohaustorius perkeus, new species
FIGURES 28, 29
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 comprising only thin setae. Armament
row on article 4 of antenna 2 even. Accessory
flagellum not reduced. Main flagellar articles of
male antenna 1 not thickened, aesthetascs 2-3
per article and occasionally present on article 1;
article 5 of male not bulbous. Coxa 1 not larger
than coxa 2; coxa 1 densely, coxa 2 sparsely
setose. Spines on dactyl of pereopod 5 set singly
in tandem. Dactyls of pereopods 6-7 vestigial.
Article 2 of pereopod 7 with widely spread pos-
terior setae. Male sternite 6 lacking ventral tooth.
Epimeron 2 bearing large tooth, with 5-6 facial
setae in vertical row(s). Tooth of epimeron 3
large. Inner rami of uropods 1-2 much shorter
than outer rami. Dorsolateral margin of peduncle
on uropod 2 with apical spines. Apicolateral cor-
ner of peduncle on uropod 3 with stout spines;
outer ramus with only setae laterally on article 1.
Telson distinctly cleft, forming slit (apical setae
not diagnostic).
DESCRIPTION OF FEMALE.?Holotype female
"f," 3.85 mm (Victoria): Head about 67% as
long as wide, rostrum about 12% as long as
remainder of head, narrow and sharp. Primary
flagellum of antenna 1 with 8 articles, articles 2-4
with 2 aesthetascs each, articles 5-7 with 1 aes-
thetasc each, accessory flagellum 6-articulate. Fa-
cial formula of setae on article 1 of antenna 1 =
4 plumes (long to short) + 2 penicillates; on
article 4 of antenna 2 = P, PPEE-P5E-P7E-7E
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 28.?Urohaustonus perkeus, new species, holotype. female "f," 3.85 mm (m = male "m,
3.42 mm).
NUMBER 360 87
mW
FIGURE 29.?Urohauslonus perkeus, new species, holotype, female "f," 3.85 mm (a ? female "a,
5.20 mm; m = male "m," 3.42 mm).
or PE, PEE-P5E-P6E-8E, article 5 clavate,
about 66% as wide as long (distal measurements),
about 58% as long as article 4, facial spine-setae
= 16 and 15 plus 6 submarginally at apex almost
continuous with dorsal armaments; article 3 with
3 facial setae; flagellum with 10 articles. Right
incisor simple, left with 2 teeth, choppers on
molars with 4 cusps; formula on right palp article
3 ? 10-1-3-1, on left = 9-1-2-2 (apparently
aberrant). Inner plate of maxilla 1 with 1 short
apical seta, 1 large apicomedial seta; outer plate
with 11 spines. Inner plate of maxilla 2 with 7
setae in medial row. Inner plate of maxilliped
with 7 medial setae, outer plate with 1 apicolat-
eral seta; palp article 2 with 8 setae in facial row
plus 1 apical facial seta, article 3 with 2 setae in
facial row. Coxa 1 subrectangular; coxa 2 similar
but narrower and slightly tapering, coxa I densely
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
setulose ventrally, about 9 of these setules elon-
gate, coxa 2 weakly setulate ventroposteriorly,
posterior margin with 3 long thin setae; coxa 3
with 20 ventral setae, mostly in posterior half,
posterior margin with 4 setae; coxa 4 with 6
ventral and 13 posterior setae, all short. Gill of
coxa 5 much larger than posterior lobe.
Setal and spine formulas on pereopod 3 = 5, 9,
2-1-1-1, 6 + 4 + 1 ; pereopod 4 = 5, 8, 2-1, 6 +
4 + 1 , posterior margin of articles 5-6 with row(s)
of medium serrations. Article 2 of pereopods 5-7
equally setose posteriorly in medium densities;
article 2 of pereopod 5 lacking mediofacial brush,
bearing numerous anterior prickles, dactyl with
6 spines in tandem.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1, segmental formulas =
20-14, 19-13, 18-13, basal setal formulas =
11-2-2-3, 7-2-2-2, 9-0-2-3.
Epimeron 2 with 5 setae in vertical row and 1
seta on ventral row.
Lateral margin on peduncle of uropod 1 with
4 setae, apicolateral corner with 1 setule; of uro-
pod 2 with pair of spines and setule at apicolateral
corner; dorsal setae = 3 and 2; setae of rami =
8-3 and 7-3, inner rami both extending 50%
along outer. Apicolateral corner of peduncle on
uropod 3 with 3 spines, medial margin with 2
setae in tandem and group of 3, ventromedial
apical row(s), with 11 mixed setae; spine formula
on lateral margin of article 1 on outer ramus =
Ee-Ee/Ee-Ee.
Telson about 1.15 times as wide as long, cleft
about halfway, setal formula = 2P, P, 1/2P, P, 1.
Lateral gland clusters well developed in pleon-
ites 4-6, more weakly developed in peduncle of
uropod 3.
Giant female "a," 5.20 mm (New South
Wales): Primary flagellum of antenna 1 with 9
articles, accessory flagellum with 8 articles, ar-
mament row on article 1 of peduncle with 5
plumes and 2 penicillates. Facial armament for-
mula on article 4 of antenna 2 =
PE-3EP-P5E-E-PE X 17; article 5 = E X 18
plus 5 apical; article 3 with 9 facial spine-setae;
flagellum with 11 articles. Right molarial chopper
with 6 cusps; right palp 3 formula = 11-1-3-1.
Coxa 1 with 15 setae in anterior long setal group,
11 in posterior short setal group, posterior margin
with 1 seta; coxa 2 with 10 ventral setules, 7
posterior setae (2 ventralmost short); coxa 3 with
28 long ventral setae and 5 posterior setae; coxa
4 with 9 ventral and 31 posterior setae. Spine and
setal formulas on pereopod 3 = 6, 10, 2-1-1-1-1,
6 + 4 + 1; on pereopod 4 = 6, 8, 2-1-1-1, 6 + 4
+ 1. Dactyl of pereopod 5 with 9 spines in tan-
dem. Epimeron 2 with 5 setae along posterior
margin and row of 4 in vertical facial row. Pe-
duncle of uropod 1 with 6 lateral setae, of uropod
2 with 5 apicolateral spines (illus.), dorsals = 6
and 7, setae of rami = 10-5 and 9-3. Uropod 3
illustrated. Telsonic setal formula = 2P, 1, P, 1
on right side, but left side like holotype. Lateral
glands are present in pleonite 3, otherwise like
holotype.
Female "g," 4.26 mm (Queensland): Primary
flagellum of antenna 1 with 8 articles, accessory
flagellum with 6 articles. Facial setal formula of
article 4 on antenna 2 = 1-3-6-13; flagellum 9-
articulate. Dactyl of pereopod 5 with 7 spines in
tandem. Epimeron 2 with 5 setae, ventralmost
small and offset anteriad. Peduncle of uropod 1
with 4 lateral and 3 dorsal setae; rami with 7 and
3 setae. Peduncle of uropod 3 like female "a"
apicolaterally; article 1 of outer ramus with 3
lateral sets of armaments.
DESCRIPTION OF MALE (male "m," 3.42 mm,
Victoria).?Generally like female with following
variations: Flagella of antenna 1 with 7 and 6
articles. Article 3 of antenna 2 with 1 plumose
and 3 simple facial setae; facial formula of article
4 = PE-PEE-P4E-PE X 12; article 5 ovoid,
about 65% as wide as long, 60% as long as article
4, facial formula = 16E and 8E apically; flagel-
lum with 10 articles. Formulas of right and left
mandibular palp article 3 = 10-1-3-1 and
11-1-3-1. Inner plate of maxilla 2 with 7 setae in
medial row. Article 2 of maxillipedal palp with 8
setae in facial row, 3 of these tightly clumped
proximally. Anteroventral edge of coxa 1 slightly
excavate. Spine and setal formulas of pereopod 3
= 4, 8, 2-1-1-1, 6 + 4 + 1 ; of pereopod 4 = 4, 7,
NUMBER 360 89
2-1, 5 + 4 + 1. Epimeron 2 with 5 setae in
vertical row and 1 seta on posterior margin. Pe-
duncle of uropod 1 with 5 lateral setae, of uropod
2 like holotype; setae of rami = 6-2 and 7-3;
inner ramus of uropod 3 slightly more elongate
than in female.
HOLOTYPE.?NMV, female "f," 3.85 mm (il-
lus.).
TYPE-LOCALITY.?CPBS 24N/872, Western
Port Bay, Victoria, Crib Point resample survey,
Aug 1972, 9 m, sand.
VOUCHER MATERIAL.?Type-locality: male
"m," 3.42 mm (illus.); male "h," 2.99 mm; AM
P.23297: female "a," 5.20 mm (illus.); AM
P.233OO: male "p;" QM W.6386:female "g," 4.26
mm.
RELATIONSHIP.?The deep cleft of the telson
suggests comparison of the species is best made to
U. pulcus, the model of the genus. Urohaustonus
perkeus differs from that model in the alate telson,
larger teeth of epimera 1-2, vertical alignment of
setae on epimeron 2, the rectangular and setose
coxae 1-2, the shortened setae of coxa 4, the weak
dactyls of pereopods 6-7, the absence of thick
spines in the armament rows of the antennae;
and more minor characters, including the pres-
ence of anterior prickles on article 2 of pereopod
5, the lack of proximolateral spines on the pedun-
cle of uropod 2 (apical only), the better posterior
setation on article 2 of pereopods 6-7, the lack of
a dactylar tooth on pereopods 3-4, and the
broader spread of facial setae on article 4 of
pereopods 3-4.
MATERIAL.?CPBS, 3 samples from 1 station
(8); WPBES, 1 sample (3); PPBES, 20 samples
from 10 stations (136); Bass Strait, 1 sample (7);
AM HRDWB, 21 samples (27); QM 5 samples
(56).
DISTRIBUTION.?Victoria, Western Port Bay to
Moreton Bay, Queensland, open sea and embay-
ment, 2-26 m, fine and coarse sand, silty sand.
Urohaustorius halci Sheard
FIGURES 30-33 (part)
Urohaustonus halei Sheard, 1936:446-449, fig. 1.
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 bearing only thin setae. Armament
row on article 4 of antenna 2 even. Accessory
flagellum not reduced. Main flagellar articles of
male antenna 1 not thickened, aesthetascs 2 per
article but usually absent on article 1; article 5 of
male antenna 2 not bulbous. Coxa 1 not larger
than coxa 2, both bearing only tiny setules. Spines
on dactyl of pereopod 5 usually set in pairs on at
least 1 locus. Dactyls of pereopods 6-7 vestigial.
Article 2 of pereopod 7 with several medium
proximoposterior setae. Male sternite 6 with long,
narrow, acute, posteriorly directed, midventral
tooth. Epimeron 2 lacking or bearing small cusp,
with 4-9 facial setae in 1-2 vertical rows. Tooth
of epimeron 3 small to large. Inner rami of uro-
pods 1-2 scarcely shorter than outer rami. Dor-
solateral margin of peduncle on uropod 2 without
spines. Apicolateral corner on peduncle of uropod
3 with thin setae, outer ramus with only setae
laterally on article 1. Telson entire, bearing both
penicillate and nonpenicillate setae.
ADDITIONAL CHARACTER.?Proximalmost seta
in row on article 1 of antenna 1 not plumose.
ADDITIONAL DIAGNOSIS DIFFERENTIATING THIS
SPECIES FROM U. metungi.?Apex of coxa 1 twisted,
thus giving subquadrate appearance from medial
view. Coxae 3-4 narrower than in U. metungi,
posteroventral apex of coxa 3 sinuous, posterior
margin of coxa 4 sinuous. Article 6 of pereopod
5 with supernumerary spine group posteriorly
(commonly 4 posterior and 3 anterior groups).
Article 4 of pereopod 6 lacking setae distally
(small gap without setae). Posterodorsal cusps of
urosomites 1-2 more pointed than in U. metungi,
with weak seta apically. Outer ramus of uropod
2 usually with lateral marginal setae or setules.
Cuticle of telson lacking large scattered setules
(at least any setules present few and small).
DESCRIPTION OF MALE.?Paratype male "e,"
3.55 mm (South Australia): Head about lb% as
long as wide, rostrum about 10% as long as
remainder of head. Primary flagellum of antenna
1 with 7 articles, articles 2-4 with 2 aesthetascs
each, articles 5-6 with 1 aesthetasc each; acces-
sory flagellum 6-articulate. Facial formula of se-
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 30.?Urohaustorius halei Sheard, paratype, male "e," 3.55 mm (j = paratype male " j , "
3.38 mm).
NUMBER 360 91
FIGURE 31.?Urohaustorius halei Sheard, paratype, male "e," 3.55 mm (j = paratype male " j , "
3.38 mm; p = paratype female "p," 3.17 mm).
92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
\v
FIGURE 32.?Urohaustonus halei Sheard, paratype, male "e," 3.55 mm (a = female "a," 5.52 mm;
j = paratype male " j , " 3.38 mm; n = juvenile "n," 2.37 mm; p = paratype female "p," 3.17
mm; Z = gill).
NUMBER 360 93
pW2
FIGURE 33. ? Upper, I'rohaustonus metungi Fearn-Wannan, male "t." 4.21 mm (g = female "gg. '
4.03 mm; j = juvenile " j , " 2.48 mm; m = juvenile "mm," 2.32 mm; q = paratype male "q,
3.87 mm). Lower, i'rohauslortus halei Sheard, p = male "p, 6.73 mm.
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
tae on article 1 of antenna 1 = 6 + 1; on article
4 of antenna 2 = 10-7-15, on article 5 = 18;
article 3 with 3 facial setae; flagellum with 8
articles. Right and left incisors simple, right laci-
nia mobilis trifid, left simple, choppers on molars
with their several cusps welded together; formula
on right and left palp article 3 = 7-1-3-1. Gland
cones on lower lip absent. Inner plate of maxilla
1 with 1 short apical seta, 1 large apicomedial
seta; outer plate with 11 spines. Inner plate of
maxilla 2 with 6 setae in medial row. Inner plate
of maxilliped with 7 medial setae, outer plate
with 1 apicolateral seta; palp of article 2 with 7
setae in facial row, article 3 with 2 setae in facial
row.
Coxa 1 broadly subtriangular, almost slipper-
shaped; coxa 2 similar but apex not as attenuate;
coxa 3 with 15 ventral setae mostly in posterior
half, posterior margin naked; coxa 4 with 4 ven-
tral and 11 posterior setae.
Setal and spine formulas on pereopod 3 = 3, 6,
2-1-1-1, 6 + 3 + 1; on pereopod 4 = 4, 6, 2-1-1,
5 + 4 + 1; posterior margin of article 5 with
serrations of mixed sizes. Article 2 of pereopods
5-7 progressively with medium to sparse setal
densities posteriorly; article 2 of pereopod 5 lack-
ing mediofacial brush, dactyl with distal to prox-
imal spine formula groups of 2-1-2-2-3-1.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental formulas =
20-14, 18-13, 17-14; basal setal formulas =
U_3_l_55 6-1-3-3, 7-6-3-4.
Epimeron lacking tooth, with 4 setae in vertical
posterior row; tooth of epimeron 3 small.
Lateral margin of peduncle on uropod 1 with
6 setae, extraordinary gap between ultimate and
penultimate setae, apicolateral corner with 1 se-
tule; of uropod 2 with only 1 apicolateral setule;
dorsal setae = 3 and 3; setae of rami = 7-6 and
6-5, outer ramus of uropod 2 with 2 lateral setae
additionally; inner rami extending 100% and 90%
along outer. Apicolateral corner of peduncle on
uropod 3 with 3 setae, medial margin naked,
ventromedial apical row(s) with 7-8 setae; spine
formula on lateral margin of article 1 on outer
ramus = ee-e-Ee-Ee.
Telson about 1.1 times as wide as long, entire,
setal formula = P, P, 1, P, 1.
Lateral and telsonic gland clusters not appar-
ent (see female "a" for probable typical condi-
tion).
Paratype male " j , " 3.38 mm (South
Australia): Article 1 of antenna 1 with 7 setae
in armament row, primary flagellum with 7 ar-
ticles, accessory with 6. Formula in armament
row on article 4 of antenna 2 = 6-5-16 (illus.),
callus at joint between articles 4 and 5 not as
strong as in paratype "e." Formula on right and
left mandibular palp article 3 = 6-1-3-2; left
lacinia mobilis also simple. Spine and setal for-
mulas on pereopod 3 = 4, 5, 2-1 - 1 , 6 + 3 + 1; on
pereopod 4 = 4, 5, 2-1-1, 5 + 3 + 1. Right dactyl
of pereopod 5 like male "e" but formula on left
side = 2-1-2-2-2-1 (illus.). Epimeron 2 with 4
setae; tooth of epimeron 3 larger than paratype
"e" (illus.). Peduncle of uropod 1 with 6 lateral
setae (illus.), rami with 7 and 6 setae; outer ramus
of uropod 2 with only 1 lateral seta. Telsonic setal
formula like male "e" but telson less elongate.
Pereonal sternite 6 with midventral hook.
Large male "g," 3.75 mm (Tasmania): Mouth-
parts not analyzed; Antenna 1 like male "e."
Formula on article 4 of antenna 2 = 2-1-7-6-22;
on article 5 = 20; medial surface of article 5
bearing fingerprintlike ridges on middle and ap-
ical calluses. Formula on article 6 of both pereo-
pods 3-4 = 6 + 4 + 1. Formula on dactyl of
pereopod 5 = 2-1-2-2-2-1-1. Setal formulas on
rami of uropods 1-2 = 7-6 and 6-5.
Large male "w," 5.23 mm (New South
Wales): Formula on dactyl of pereopod 5 =
3-2-2-2-3-2-1. Naked apicoposterior margin on
article 4 of pereopod 6, typical of this species,
longer than in illustration. Epimeron 2 with 5
setae (dorsalmost small); epimeron 3 with very
long tooth as in E. metungi (like female "a," illus.).
Setal formula on rami of uropods 1-2 = 9-7 and
8-7, outer ramus of uropod 2 with 4 lateral setae
additionally; peduncle of uropod 1 with 6 lateral
setae lacking extraordinary gap; of uropod 2 with
2 apicolateral setules; of uropod 3 with 4 apico-
NUMBER 360 95
lateral setae. Telsonic setal formula = P, P, 1, 1,
P, 1.
Small male "r," 3.45 mm (Pambula Beach,
NSW): About the size (3.45) of South Austra-
lian paratypes, with cusp of epimeron 3 better
developed, though neither as long nor as straight
as that on the very large specimens from New
South Wales and Queensland ("a," illus.). Tel-
sonic formula = P, P, 1, 1, P, 1.
Large male "pp," 6.73 mm (largest known
specimen; Queensland): Armament formulas
on article 1 of antenna 1 = 1 2 (no seta plumose);
on article 4 of antenna 2 = 17-12-34; primary
flagellum of antenna 1 with 10 articles, accessory
with 9, flagellum of antenna 2 with 11 articles.
Right mandibular incisor with 4 teeth, rakers
unusual (illus.), formulas on right palp article 3
= 9-1-3-1, left = 7-1-3-1. Medial row on inner
plate of maxilla 2 with 8 setae. Palp article 2 of
maxilliped with 9 setae in facial row, article 3
with 2 and 1 additional apicomedial seta. Coxa
1 illustrated; coxa 2 also sharp. Spine and setal
formulas on pereopod 3 = 4, 8, 2-1-1-1-1, 7 + 4
+ 1, on pereopod 4 = 4, 8, 2-1-1-1-1-1, 7 + 4
+ 1, posterior margin of article 4 with large tooth
scales as on article 5 of younger specimens. Article
6 of pereopod 5 with 3 supernumary posterior
spine groups instead of 1; formula on dactyl of
pereopod 5 = 1-2-2-3-3-3-3 (illus.), also with
marginal serrations on apex (illus.). Article 4 of
pereopod 6 with large gap between last posterior
seta and apex. Epimeron 2 with square postero-
ventral corner, face with 11 setae in 2 rows (illus.).
Tooth of epimeron 3 large. Peduncle of uropod 1
with 8 lateral setae, extraordinary gap scarcely
present, peduncle of uropod 2 with 1 apicolateral
setule, dorsal setae = 4 and 1, setae of rami = 9-9
and 8-8, outer ramus of uropod 2 also with 3
lateral setae additionally. Peduncle of uropod 3
with 5 apicolateral setae and 3 large medial setae
in tandem, otherwise see illustration (large medial
setae omitted). Telson elongate (illus.) formula
= 2P, 1,P, 1, 1.
DESCRIPTION OF FEMALE.?Paratype female
"p," 3.17 mm (South Australia): Generally like
male "e" with following variations: Armament
row of article 1 on antenna 1 = 5; of article 4 on
antenna 2 = 2-6-5-17; on article 5 = 14. Primary
flagellum of antenna 1 with 7 articles, accessory
with 6 articles; flagellum of antenna 2 with 8
articles; article 4 of peduncle scarcely calloused.
Mouthparts not examined. Spine and setal for-
mulas on pereopod 3 = 4, 5, 2-1-1, 5 + 4 + 1; on
pereopod 4 = 4, 4, 2-1-1, 5 + 3 + 1. Brood plates
normal, setose. Dactyl of pereopod 5 like male
"e" but proximal spine shorter. Epimeron 3 like
male " j " (elongate form), but ventral setule ab-
sent. Outer ramus of uropod 2 with 1 lateral seta
at M. 75. Medium setal formula on lateral margin
of article 1 on outer ramus of uropod 3 = e-Ee-Ee
(illus.); ventral peduncular setae 6-7, none offset.
Telson like male " j " but setae symmetrical.
Giant female "a," 5.52 mm (Queensland): For-
mula on dactyl of pereopod 5 = 3-2-2-2-2-2.
Epimeron 2 with slightly produced posteroventral
corner, with 9 setae in 2 rows (illus.); epimeron 3
with extremely elongate tooth (illus.). Peduncle
of uropod 1 with 8 lateral setae lacking extraor-
dinary gap, of uropod 2 with 1 apicolateral setule;
dorsal setae = 3 and 1, setae of rami = 9-8 and
8-6, outer ramus of uropod 2 with 1 lateral seta
additionally. Apicolateral corner of peduncle on
uropod 3 with 3 setae. Telson unusually truncate
apically, setal formula = P, P, 1, 1, P, 1, with
inner glands with U. metungi.
Female "m," 4.12 mm (Queensland): Formula
on dactyl of pereopod 5 = 3-2-3-1. Epimeron 2
with rounded posteroventral corner, with 9 setae
in 2 cycles. Tooth of epimeron 3 elongate. Outer
ramus of uropod 2 with 2 lateral setae. Brood
plates with vestigial setae but internally showing
next instar with normal setae.
Giant female "b," 5.72 mm (Queensland, spec-
imen subsequently lost in transit): Outer ramus
of uropod 2 with only 1 lateral seta positioned
very apicad. Apicolateral corner of peduncle on
uropod 3 with only 1 apicolateral seta.
DESCRIPTION OF JUVENILE (juvenile ikn," 2.37
mm; Queensland).?Armament row on article 1
of antenna 1 = 3 long; primary and accessory
flagella each with 6 articles. Spine and setal
formulas of both pereopods 3-4 = 2, 4, 2-1-1, 5
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
+ 3 + 1 . Dactyl of left pereopod 5 with 3 spines
in tandem; of right side with formula of 1-1-2-1
(illus.).
Epimeron 2 with 2 setae; epimeron 3 well
produced as illustrated for male " j . " Peduncle of
uropod 1 with 4 lateral setae, uropod 2 with 1
apicolateral setule, dorsal setae = 2 and 1, setae
of rami = 5-4 and 4-4. Apicolateral corner of
peduncle on uropod 3 with 1-2 setae. Telsonic
formulas, right = P, P, 1, 1, P, left = P, P, P, 1.
TASMANIAN SPECIMENS.?Conforming well to
paratypes of South Australia.
VICTORIAN SPECIMENS.?Conforming to para-
types.
NSW SPECIMENS.?Differing from those of
South Australia in (1) much larger size (up to 5.0
mm) compared with 3.0+ mm; (2) much blunter
and in largest specimens more heavily calcified
coxa 1 with apex forming scoop; (3) narrower
coxa 3 with distinctly sinuous apex; (4) leaf-
shaped coxa 4, subtly distinct from paratypes; (5)
more strongly setose lateral margin of outer ramus
on uropod 2, with setae extending almost the
entire length of the ramus; (6) more robust dactyls
of pereopods 6-7; (7) more numerous setae else-
where; (8) longer and more acute tooth of epi-
meron 3; (9) heavy armament of telson, asym-
metrical in juveniles, symmetrical in larger adults.
QUEENSLAND SPECIMENS.?Conforming to U.
halei diagnosis, except for enlarged tooth of epi-
meron 3, bridging gap between metungi and halei;
much larger than paratypes and with (1) broader
upper lip (also true of NSW); (2) broader, blunter
coxae 2-4; (3) more robust dactyl of pereopods
6-7; (4) broader article 5 of pereopod 7; (5) weak
beginning of division of setae into 2 groups; (6)
longer, less upturned cusp on epimeron 3, with
broader flatter sinus, thus closest to U. metungi.
LECTOTYPE.?SAM, originally designated in
print as "type male 2.5 mm." Allotype female
(C.2081) originally designated in print as
"ovigerous type female, 1.9 mm."
TYPE-LOCALITY.?Sellick's Reef, Gulf of St.
Vincent, South Australia, burrowing in sand on
sandy patches on reef, M M . Hale, Apl. (sic)
1936.
VOUCHER MATERIAL.?Paratypes (presumably
from type-locality, newly labeled by us): male
"e," 3.55 mm (illus.); male " j , " 3.38 mm (illus.);
female "p," 3.17 mm (illus.); female "q," 2.90
mm (measured by parabolic rather than straight-
line method); Tasmania, Kingston Beach, 6 Feb
1975, T.M. Walker: male "g," 3.75 mm; MMD,
Pambula Beach, NSW: male "r," 3.45 mm; AM
P.23343: male "u," 5.04 mm; male "w," 5.23
mm; QM W.6386: female "a," 5.52 mm (illus.);
female "m," 4.12 mm; juvenile "n," 2.37 mm
(illus.); QM W.6384: female "b," 5.72 mm; QM
W.6344: male "pp," 6.73 mm (illus.); QM 5508:
male "nn" 4.91 mm.
RELATIONSHIP.?This species, together with U.
metungi and U. gunni, form a group characterized
by elongate inner rami of uropods 1-2, rounded
epimeron 2 (except in huge seniles, and Queens-
land specimens), loss of tooth on dactyls of per-
eopods 3-4, thinness of armaments on antennae
1-2 and on apicolateral corner of peduncle on
uropod 3, uncleft telson, poorly setose coxae 1-2
and lack of lateral spination on the peduncle of
uropod 2, lack of lateral urosomal glands but the
presence of glands in the telson. Urohaustorius gunni
is easily recognized by the unusual, enlarged, and
hooked coxa 1 and the spikes on pereonite 7 and
coxa 7; however, U. halei and U. metungi are very
difficult to segregate.
Generally U. halei differs from U. metungi in the
doubling of spines on the dactyl of pereopod 5
and in the shorter tooth of epimeron 3. Large
adults of U. halei ohen have the tooth of epimeron
3 almost as long as that of adults and juveniles of
U. metungi. Presumably juveniles of U. halei (figure
32 "n", herein, from Queensland) have unpaired
spines on the dactyl of pereopod 5. Juveniles from
Queensland are almost as large as certain adults
from Victoria, so that the distinctions will require
an understanding of the instar sequences in var-
ious geographic areas. (See "Relationship" of U.
metungi, page 102.)
Both species show variability in a number of
characters for which only general discussion is
needed. For example, the number of setae in-
creases with body size in both species. Males of
NUMBER 360 97
both species have narrower dactyls on pereopod
5 than do females. Precise telsonic shapes vary
from sample to sample and within samples.
MATERIAL.?South Australia, St. Vincent's
Gulf, in sand on reef (holotype and many para-
types); Victoria, MMD samples, Back Beach, San
Remo, Western Port (many), Wilson's Promon-
tory, mouth of Tidal River (many); Tasmania,
Walker sample, Kingston Beach, South end (2);
New South Wales, HDWBS, 11 samples (22), 1
of these samples at Stockton Beach, 6 m, Nov
1975, 10 samples from 500 m E of Belmont Beach
(offshore), 16 km S of Hunter River, 12-15 m,
various transects, Aug 1975 to Feb 1976; MMD
sample, Pambula surf beach (1); Queensland, 5
samples (42), 3 of these samples from Middle
Banks, Moreton Bay, 1 from Moon Creek on
Fraser Island and 1 from mouth of Noosa River,
inside Noosa Bar, 12-15 m.
DISTRIBUTION.?South Australia, St. Vincent's
Gulf to Moreton Bay, Queensland, littoral to 15
m, sand, sand and mud.
Urohaustorius vercoi Sheard
Urohaustorium vercoi Sheard, 1936:449, fig. 1 (part).
REMARKS.?The only specimen of this species
we have found labeled as such in the South
Australian Museum is called an "allotype" but it
clearly is not the species Sheard described as U.
vercoi; indeed it is not even in the genus Urohaus-
torius.
We have also looked in the Australian Mu-
seum, the Queensland Museum, the National
Museum of Victoria, and the Western Australian
Museum for Sheard's types, to no avail. Sheard's
general amphipod materials became scattered
even before his death, as shown by the fact that
some of his specimens from South Australia have
been found in the Australian Museum and the
Western Australian Museum where he either sent
or personally deposited them.
Urohaustorius vercoi is based on a skimpy descrip-
tion and figures, not enough to confirm its iden-
tity; however, the short inner rami of uropods
1-2, coupled with the elongate article 2 ofgnath-
opod 2 and, we infer from Sheard's comparison
to U. halei, an uncleft telson (or almost so if he
did not notice a tiny invagination), suggests to us
that the species lies near our new species, U.
pentinus. However, Sheard (1936) noted or de-
picted the relative smallness of coxa 3 in compar-
ison to coxa 4, and the strongly produced coxa 4
that would appear to be firm distinctions. The
coxal sizes suggest our new genus and species,
Narunius tallerkus, but the shape of coxa 4 and the
proportions of rami of uropods 1-2 are not con-
sistent with our species.
Urohaustorius metungi Fearn-Wannan
Figures 33 (part)-35
Urohaustorius metungi Fearn-Wannan, 1968:31-37, parts of
figs. 1-3.
Diagnosis.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 bearing only thin setae. Armament
row on article 4 of antenna 2 even. Accessory
flagellum not reduced. Main flagellar articles of
male antenna 1 not thickened, aesthetascs 2 per
article but usually absent on article 1, article 5 of
male antenna 2 not bulbous. Coxa 1 not larger
than coxa 2; both bearing only tiny setules. Spines
on dactyl of pereopod 5 (usually) set singly in
tandem. Dactyls of pereopods 6-7 vestigial. Ar-
ticle 2 of pereopod 7 with long posterodorsal setae
and setules. Male sternites 5-6 each with ventral
tooth. Epimeron 2 lacking cusp, commonly with
4 facial setae in vertical row. Tooth cusp of
epimeron 3 huge. Inner rami of uropods 1-2
scarcely shorter than outer rami. Dorsolateral
margin of peduncle on uropod 2 without spines.
Apicolateral corner of peduncle on uropod 3 with
thin seta(e); outer ramus with only setae laterally
on article 1. Telson entire, bearing only penicil-
late setae.
ADDITIONAL CHARACTER.?Proximalmost seta
in row on article 1 of antenna 1 plumose.
ADDITIONAL DIAGNOSIS DIFFERENTIATING THIS
SPECIES FROM U. halei.?Apex of coxa 1 scarcely
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 34.? Urohaustonus metungi Fearn-Wannan, male "t," 4.21 mm (u = male "u," 4.49 mm;
w = female "w," 3.82 mm; Z = male sternites).
NUMBER 360 99
HYI ^-^ \yy Y \ ^^r=?
FIGURE 35.?Urohaustorius metungi Fearn-Wannan, male "t," 4.21 mm (Y = pleopod; Z = gill).
100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
twisted or not. Coxae 3-4 broader than in U.
halei, posteroventral apex of coxa 3 broad, not
sinuous; posterior margin of coxa 4 concave; an-
terodistal corner of coxa 3 forming marked heel.
Article 6 of pereopod 5 without supernumerary
spine group posteriorly (commonly with 3 poste-
rior and 3 anterior groups). Article 4 of pereopod
6 bearing setae continuously to apex. Posterodor-
sal cusps of urosomites 1-2 blunter than in U.
halei and unarmed. Outer ramus of uropod 2
naked laterally. Cuticle of telson with scattered
large setules varying from few in Victorian and
most NSW specimens to many in some specimens
from Queensland.
DESCRIPTION OF MALE.?Male "t," 4.21 mm
(Mallacoota): Head 70% as long as wide, ros-
trum about 10% as long as remainder of head.
Primary flagellum of antenna 1 with 8 articles,
articles 2, 6-7 each with 1 aesthetasc, articles 3-
5 with 2 each, accessory flagellum 6-articulate.
Facial formula of setae on article 1 of antenna 1
= 1 plume and 9 other mixed setae; on article 4
of antenna 2 = 8-7-22; on article 5 = 16; article
3 with 5 facial setae; flagellum with 8 articles.
Right and left incisors simple, right lacinia mob-
ilis bifid, left simple, choppers on molars with
their several cusps welded together, forming a
circlet of 13 badly worn cusps, formula on right
palp article 3 = 9-1 -4 -1 , on left = 9-1-3-1. Inner
plate of maxilla 1 with 1 short apical seta, 1 large
apicomedial seta; outer plate with 11 spines. In-
ner plate of maxilla 2 with 11 setae in medial
row. Inner plate of maxilliped with 7 medial
setae, outer plate with 1 apicolateral seta; palp
article 2 with 7 setae in facial row, article 3 with
2 setae in facial row.
Coxa 1 broadly subtriangular; coxa 2 similar,
apex somewhat more attenuate but not as sharp;
coxa 3 with 19 ventral setae mostly in posterior
half, posterior margin naked; coxa 4 with 3 ven-
tral and 6 posterior setae.
Setal and spine formulas on pereopod 3 = 6, 6,
2-1-1-1-1, 7 + 4 + 1; on pereopod 4 = 5, 6, 2-
1-1-1-1,6 + 4 + 1; posterior margin of article 5
with interrupted groups and double groups of
serrations. Article 2 of pereopods 5-7 progres-
sively with medium to sparse setal densities pos-
teriorly; article 2 of pereopod 5 with mediofacial
brush lacking or weakly developed, dactyl with 5
spines in tandem.
Pereonal sternites 5-6 each with midventral
process.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 1, 2 and 1; segmental formulas =
22-16, 20-15, 18-17 (inner proliferate); basal
setal formulas = 10-2-4-4, 9-1-4-4, 10-2-4-5.
Epimeron 2 lacking tooth, with 4 setae in ver-
tical posterior row; tooth of epimeron 3 huge.
Lateral margin of peduncle on uropod 1 with
5 setae, extraordinary gap between ultimate and
penultimate setae, apicolateral corner with one
setule, of uropod 2 with only 1 apicolateral setule;
dorsal setae = 3-4 and 2; setae of rami = 7-5
and 6-6; outer ramus of uropod 2 naked, inner
rami extending 90% and 85% along outer. Api-
colateral corner of peduncle on uropod 3 with 1
setule, medial margin with 2 long setae, ventro-
medial apical rows with 11 setae; spine formula
on lateral margin of article 1 on outer ramus = 0,
short setae = 1-1-0, medium setae = 0-1-1, long
setae = 0-0-1.
Telson about 1.1 times as wide as long, entire,
setal formula = 2P, P/2P, P.
Lateral gland clusters absent; telson with cen-
tral gland cluster.
Paratype male "q," 3.87 mm (our designation
from paratype lot, Lake King): Armament for-
mula on article 1 of antenna 1 = 5; on article 4
of antenna 2 = 2-7-4-14; article 5 = 1 1 ; article
3 = 5. Formula on right and left mandibular palp
article 3 = 7-1-3-1. Spine and setal formulas on
pereopod 3 = 5, 6, 2-1-1, 5 + 3 + 1; same on
pereopod 4 for last 2 numbersets. Dactyl of per-
eopod 5 with 4 spines. Peduncle of uropod 1 with
4 lateral setae; setae on rami of uropods 1-2 =
6-4 and 5-5. Inner ramus of uropod 3 with only
1 lateral and 3 medial setae, outer ramus lateral
formula, short setae = e-Ee. Specimen like male
"t" in other characters. Maxillipeds missing, pleo-
pods and cuticle not examined, some spines on
outer plate of maxilla I with more than 2 cusps.
NUMBER 1560 101
Fearn-Wannan mentioned large eyes in male;
paratype "q" shows eye capsules but no pigment.
Male "u," 4.49 mm (Mallacoota): Slightly
better developed than male "t"; formula on right
mandibular palp article 3 = 7-1-3-1. Formula
on article 6 of pereopod 3 = 6 + 4 + 1; on
pereopod 4 = 5 + 4 + 1. Epimeron 2 with 5 setae.
Peduncle of uropods 1-2 each with 3 dorsal setae;
setae on rami = 8-6 and 7-5.
Male "f," 4.39 mm (Tuross River, NSW):
Like Victorian U. metungi, with 4 spines in tandem
on dactyl of pereopod 5 and with elongate tooth
on epimeron 3; dactyl of pereopod 5, however,
not inflated.
DESCRIPTION OF FEMALE.?Female "w," 3.82
mm (Mallacoota): Generally like male but with
following variations: Armament formulas on ar-
ticle 1 of antenna 1 = 6; on article 4 of antenna
2 = 5-4-13; on article 5 = 13; on article 3 = 4;
flagellum of antenna 2 with 8 articles (flagella of
antenna 1 imperfect). Spine formulas on articles
5 and 6 of both pereopods 3 and 4 = 2-1-1, 5 +
3 + 1 . Dactyl of pereopod 5 with 3 spines in
tandem (illus.). Brood plates normally setose.
Epimeron 2 with 4 setae. Setae on rami of uropods
1-2 = 6-5 and 4-4. Lateral setal formula on
article 1 of outer ramus = e-Ee. Lateral glands
weakly developed in pleonites 3-6, peduncle and
outer ramus of uropod 3, and strongly developed
in telson.
Large female "h," 5.13 mm (Merimbula):
Dactyl of pereopod 5 with 5 spines in tandem,
most proximal very short. Epimeron 2 with 5
setae.
Female "a," 3.83 mm (Queensland, Maroochy
River): Dactyl of pereopod 5 with 5 spines in
tandem plus most proximal spine well disjunct.
Female "k," 4.66 mm (Merimbula): Dactyl
of pereopod 5 thin (like U. halei), with 3 apical
spines in tandem; telson almost as long as in male
"pp" of U. halei.
Female "gg," 4.03 mm (Queensland): Smaller
spine and setal counts; coxa 3 illustrated, varia-
tion also occurring in specimens of Victorian U.
metungi and U. halei; dactyl of pereopod 5 formula
= 1-1-2-1 (illus.), thus scarcely distinct from U.
metungi; epimeron 2 with 3 setae; tooth of epime-
ron 3 large; telson very short (illus.).
Juvenile female "mm," 2.52 mm (Queens-
land): Formula on dactyl of pereopod 5 = 1 - 1
(illus.). Epimeron 2 with 2 setae; tooth of epime-
ron 3 large. Peduncle of uropod 1 with 3 lateral
setae, extraordinary gap present; dorsal setae of
uropods 1 -2=1 and 1; setae on rami = 4-3 and
3-2. Telson ordinary, shaped as in male "t" of
Victorian U. metungi (bearing elongate tooth of
epimeron 3).
DESCRIPTION OF JUVENILE (juvenile " j , " 2.48
mm, Merimbula).?Dactyl of pereopod 5 with 2
apical and 1 anterior spines, all in tandem (illus.).
Telson also elongate but apex truncate.
VARIATION IN SPINE DOUBLING.?About 10% of
adults from New South Wales and Queensland
show spine doubling on the dactyl of pereopod 5
(Table 1). Victorian specimens examined, includ-
ing the holotype and paratype, do not. The data
vaguely suggest that frequency of doubling in-
creases towards Queensland and away from Vic-
toria but is not statistically significant and must
be reexamined from larger samples than we now
have on hand.
Although in U. halei doubling occurs at 2 or
more loci and trebling may occur at 1 or more
loci, only 2 males of U. metungi (from Merimbula)
show double doubling (2 loci doubled). All other
examples are confined to a single locus, usually
the secondmost proximal, and most frequently
doubling occurs only on 1 side (either left or right
pereopod 5).
Unfortunately specimens of U. metungi with
doubled spination spoil what would otherwise be
a facile way to distinguish this species from U.
halei, at least in adult stages.
QUEENSLAND SAMPLES.?A degree of variation
in individual size of adults was observed within
samples and between samples but we have not
been able to detect any further taxonomic divi-
sions suggested by this allomorphy. Considerable
variation in precise shape and setular equipment
of the telson was also evident, shape ranging from
quite tall (Noosa River) to broad and flattened
(Serpentine Creek), though always with variation
102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Spine doubling on dactyl of pereopod 5 in
Urohaustonus metungi
Location
NSW
Merimbula
Tuross River
Georges River
Queens Lake
Queensland
Jacksons Creek
Serpentine Creek
Noosa River
Maroochy River
Ninghi Creek
Culboolture River
Totals
No. specimens
Specimens doubling
Percent doubling
les
sa
m
p
o
No
.
3
44
3
9
4
8
3
3
2
2
n
en
s
.?I
en
217
225
41
76
104
152
31
11
23
7
887
88
10
V
-D
ed
 
1 
si
do
ub
l
M
ai
15
10
4
0
3
2
4
1
3
2
de
s
ed
 
2 
si
do
ub
l
d
M
ai
0
0
1
0
0
1
2
0
0
1
sid
e 
I
ib
led
 
1
0
-a
?sid
n
Fe
m
8
9
2
0
2
0
1
0
4
1
sid
es
bl
ed
 
2
;es
 
do
u
Fe
m
1
3
4
0
0
0
1
0
2
0
-a
m
do
ub
h
tw
ic
e
M
ai
2
0
0
0
0
0
0
0
0
0
bl
ed
:e
 
do
ui
H
M
J
 S3(
Fe
m
0
0
0
0
0
0
0
0
0
0
"
ce
nt
ag
e
in
g 
pe
i
do
ub
l
?
To
u
12
10
27
0
5
2
26
9
39
57
within samples. Specimens from QM W4499
(Serpentine Creek), with extremely broad flat-
tened telson and very abundant subcuticular tel-
sonic setules, intergrade through many stages to
the Victorian holotype in which the telson is
moderately tall and rounded and its setules small
and sparse. Dactyls of pereopods 6 and 7 are more
robust than in Victorian specimens and article 5
of pereopods 5 and 6 proportionally longer. Fre-
quently the rami of uropod 2, particularly in
males, appear to be less disparate in length than
the rami of uropod 1.
TUROSS RIVER SAMPLES.?Compared with
those from Queensland, Tuross River specimens
tend to have a broader coxa 4, more heavily
setose article 4 of pereopod 6 and a more atten-
uate cusp on epimeron 3. The shape of the telson
is commonly cordate compared with the more
rounded forms from other samples.
MERIMBULA SAMPLES.?Telsons are extremely
variable in shape.
HOLOTYPE.?NMV J.151, male, length un-
known.
TYPE-LOCALITY.?Lake King, Tambo Bay.
VOUCHER MATERIAL.?Type-locality: paratype
male "q," 3.87 mm (illus.); QM W.4147: female
"gg," 4.03 mm (illus.); juvenile "mm," 2.52 mm
(illus.); AM P. 24906: male "f," 4.39 mm; Mal-
lacoota Inlet, Hutchings and Kudenov coll., male
"t," 4.21 mm (illus.); same, Kudenov, first sam-
ple: female "w," 3.82 mm (illus.); second sample:
male "u," 4.49 mm (illus.); AM, Merimbula 37B:
female "h," 5.13 mm; female "k," 4.66 mm;
juvenile " j , " 2.48 mm (illus.); QM W.4528: fe-
male "a," 3.83 mm; and specimens from QM
5504 and 5531.
RELATIONSHIP.?At first this species was differ-
entiated from U. halei by the undoubted spine sets
on the dactyl of pereopod 5 coupled with the
huge tooth of epimeron 3; but the first distinction
was found to be false in a significant number of
specimens and the second too difficult to practice
with sure results. A very careful study was made
hundreds of specimens that showed that the fol-
lowing characters reliably distinguish U. metungi
from U. halei: (1) the plumose condition of the
proximal seta in the main setal row of article 1
on antenna 1; (2) the lack of a small distal gap in
the posterior setation of article 4 on pereopod 6;
and (3) the absence of marginal nonpenicillate
setae on the telson (only penicillate setules are
present, whereas halei bears these penicillate se-
tules plus ordinary simple setae). In addition,
males of metungi have 2 blunt sternal teeth, 1 each
on pereonites 5 and 6, whereas halei has only 1
curved pointed tooth on pereonite 6.
By dividing the species on these points, the
confusion about spine doubling on the dactyl of
pereopod 5 in a small percentage of metungi ap-
pears to be clarified: no doubling was found in
type material of metungi, whereas infrequent dou-
bling was found in NSW and Queensland speci-
mens, mainly on either right or left sides and
NUMBER 360 103
usually just in the second most proximal spine
set; but on both sides together in only 16 speci-
mens.
MATERIAL.?Victoria, Lake King, holotype
and paratypes; Mallacoota, Hutchings and
Kudenov coll. (5), MMD coll. (30+). NSW, AM,
Merimbula, Day and Kudenov coll., 3 samples
(217); Tuross River, Eurobodalla Shire Estuary
Survey, 44 samples (341); Pittwater, 2 samples
(2); Wallaga Lake (1); EBS, Georges River, 5
samples (42); Queen's Lake, 9 samples (76);
Queensland Museum, 24 samples (316).
DISTRIBUTION.?Victoria, Gippsland Lakes to
Moreton Bay, Queensland, intertidal estuarine.
Urohaustorius gunni, new species
FIGURES 36, 37
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 bearing only thin setae. Armament
row on article 4 of antenna 2 even. Accessory
flagellum not reduced. Main flagellar articles of
male antenna 1 [unknown], aesthetascs [un-
known], article 5 of male antenna 2 [unknown].
Coxa 1 much larger than coxa 2, extended ven-
trally and hooked forward, both coxae bearing
only tiny setules. Spines on dactyl of pereopod 5
usually set singly in tandem, but occasionally in
pairs. Dactyls of pereopods 6-7 vestigial. Article
2 of pereopod 7 sparsely armed posteriorly. Male
sternites 5-6 [unknown]. Epimeron 2 lacking
tooth, with about 6 facial setae in vertical row.
Tooth of epimeron 3 small. Inner rami of uropods
1-2 about as long as outer rami. Dorsolateral
margin of peduncle on uropod 2 without spines.
Apicolateral corner on peduncle of uropod 3 with
thin setae, outer ramus with only setae laterally
on article 1. Telson entire [apical setae not diag-
nostic].
ADDITIONAL CHARACTERS.?Article 3 of an-
tenna 2 with dorsoventral cusp; coxa 7 with spike.
DESCRIPTION OF FEMALE.?Holotype female
"i," 5.37 mm (New South Wales): Head almost
75% as long as wide, rostrum about 18% as long
as remainder of head, sharp. Article 1 of antenna
1 with 8 (6) very fine dorsofacial setae, penicillates
absent; article 2 with 3 long distal fine setae in
addition to numerous strong plumose setae; pri-
mary flagellum 8-articulate, aesthetasc formula
= 0, 0, 1, 2, 2, 1, 1, 0; accessory flagellum 7-
articulate. Article 3 of antenna 2 with 4 facial
setae and large sharp dorsal cusp; facial formula
on article 4 = PE-P-14E-3E-8E-20E+, in sev-
eral irregular rows submarginally; article 5 about
46% as wide as long, 62% as long as article 4,
facial armament consisting of many submarginal
setae, and a slightly disjunct set of 5 similar
apicals, stout spines absent; flagellum with 9
articles. Right and left incisors weakly bifid, right
lacina mobilis bifid, left simple, choppers on mo-
lars forming only simple hump; formula on right
and left palp article 3 = 9-1-3-1. Inner plate of
maxilla 1 with 1 short apical seta, 1 large apico-
medial seta; outer plate with 11 spines, some with
3 side teeth. Inner plate of maxilla 2 with 8 setae
in medial row. Inner plate of maxilliped with 7
medial setae, outer plate with 1 apicolateral seta,
palp article 2 with 9 setae in facial row, article 3
with 2 setae in facial row.
Coxa 1 extended deeply and hooked forward;
coxa 2 broadly subrectangular; coxa 3 with 22
ventral setae mostly in posterior half, posterior
margin naked; coxa 4 with 7 ventral and 30+
posterior setae, mostly short. Apex of gnathopod
2 as illustrated for U. metungi.
Setal and spine formulas on pereopod 3 = 5, 8,
2-1-1-1-1-1, 7 + 4 + 1; on pereopod 4 = 5, 7,
2-1-1-1-1, 7 + 4 + 1, spines exceptionally
smooth; posterior margin of article 5 with even,
weakly interrupted serrations. Article 2 of pereo-
pods 5-7 progressively with medium to sparse
setal densities posteriorly, filled with leached
glandular tissue (not shown in Illustrations), ar-
ticle 2 of pereopod 5 lacking mediofacial brush,
dactyl spine formula = 1-1-3-1-1-1-1, article 2
of pereopod 7 with unusual sharp bevel postero-
ventrally, posterior margin with 1 fine proximal
seta and 2 setules. No sternal process present on
pereonite 6. Brood plates normally setose.
Peduncular spine formulas of pleopods 1-3 =
104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 36. ? I'rohaustonus gunni, new species, holotype, female "i ," 5.37 mm (j = young female
" j , " 3.66 mm; Y = sternal spike; Z = gill).
NUMBER 360 105
D5 QP5
FIGURE 37.?Urohaustonus gunni, new species, holotype, female "i," 5.37 mm.
106 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
2 and 1, 2, and 0, 2 and 0; segmental formulas
= 24-18, 21-13, 21-17; basal setal formulas =
15_3_5_45 9-3-4-2 (or 8-1-3-1), 10-2-4-4.
Epimeron 2 lacking tooth, with 6 setae in ver-
tical row directly on posterior margin; tooth of
epimeron 3 small. Peduncle of uropod 1 about
half as wide as long, subequal to or slightly longer
than outer ramus, lateral margin with 8 setae,
extraordinary gap between ultimate and penul-
timate setae, apicolateral corner with 1 setule;
peduncle of uropod 2 much shorter than outer
ramus, less than twice as wide as long, with only
1 apicolateral setule; dorsal setae = 3 and 2; setae
of rami = 9-8 (or 7) and 8-6, outer ramus of
uropod 2 with 2 lateral setae additionally; inner
rami extending 100% and 98% along outer. Api-
colateral corner of peduncle on uropod 3 with 3
setae, medial margin with 2 large setae in tandem,
ventromedial apical rows with 12 setae; setal
formula on lateral margin = ee-e-ee-Ee-Ee-Ee.
Telson about 1.15 times as wide as long, entire,
setal formula = 2P, 2, P, 1.
Lateral and telsonic gland clusters not appar-
ent.
Young female " j , " 3.66 mm (New South Wales):
Armament formulas on article 1 of antenna 1 ?
5; on article 4 of antenna 2 = P, P, P7E-3E-4E-
P7E-7E, in double row. Article 5 of antenna 2
long and ovoid, 45% as wide as long, 70% as long
as article 4, facial armament = 17E + 6E apically;
primary and accessory flagella of antenna 1 each
with 7 articles on 1 side or 8 and 6 on the other,
flagellum of antenna 2 with 9 articles. Formulas
of mandibular palps = 7-1-3-1. Spine and setal
formulas of pereopod 3 = 4, 6, 2-1-1-1-1, 7 + 3
+ 1; of pereopod 4 = 4, 6, 2 - l - l - l - ( l ) , (6-7) +
4 + 1 . Dactyl of pereopod 5 with 7 spines. Epi-
meron 2 with 4 setae. Uropod 1 with 6 lateral
setae, gap weak, dorsal setae of uropods 1-2 = 3
and 1-2, setae of rami = 6-7 and 6-6. Peduncle
of uropod 3 with only 2 apicolateral setae, ven-
trally with 8-9. Coxae, epimera, and pereopods
checked, mouthparts not studied. Telsonic for-
mula = 2P, 1, P, 1 on left side, right like holotype.
Pereonite 6 with ventral spike (illus.).
Female "h," 4.10 mm: Sternal spikes absent.
HOLOTYPE.?AM P.23342, ovigerous female
"i," 5.37 mm (illus.).
TYPE-LOCALITY.?16 km south of Hunter
River, 500 m east of Belmont Beach, New South
Wales, 33?02'36" S, 151? 40'56" E, 22 Aug 1975,
12 m, sand.
VOUCHER MATERIAL.?AM P.24926, female
"h," 4.10 mm; AM P.23335, young female " j , "
3.66 mm (illus.).
RELATIONSHIP.?This species appears to be an
advancement from the U. halei-metungi group, in
which coxa 1 has been grossly modified into a
slipper spike and coxae 6-7 plus pereonite 7 are
produced sharply posteriorly into extended alae.
Epimeron 3 is unusual in the extreme loss of
lateral definition, and the very close position of
the setae to the posterior margin. Article 3 of
antenna 2 is cuspidate dorsally. Article 2 of per-
eopod 7 is of very unusual shape.
MATERIAL.?HRDWB, 4 samples (4).
DISTRIBUTION.?New South Wales, 4-16 km
south of Hunter River, 10-15 m (about 500-1500
m offshore), sand.
Gheegerus, new genus
DIAGNOSIS.?Rostrum weak. Peduncle of an-
tenna 1 short, stout, articles 2 and 3 of peduncle
progressively shortened, weakly geniculate; pri-
mary flagellum elongate, accessory flagellum
somewhat shortened. Antenna 2 of haustorius
form, article 4 expanded, article 5 small, articles
4-5 with facial armaments, article 4 with long
ventral setae and subventral clusters of bifid setae.
Aesthetascs simple. Mandibular incisors ex-
tended, thin, poorly toothed; right and left laci-
niae mobiles not alike; rakers usually serrate, 2
on right mandible, 3 on left; molar broad and
bulky, not strongly extended, with weak tritura-
tive surface, with cuspidate accessory chopper;
setae of palp article 3 awned. Mandibular lobes
of lower lip well developed. Inner plate of maxilla
I small, slender, poorly setose, outer plate with
I1 spines; palp shortened, uniarticulate, palpar
setae attached in 2 sets. Inner plate of maxilla 2
with weakly submarginal oblique row of setae.
NUMBER 360 107
Maxillae and maxillipeds lacking baler lobes.
Outer plate of maxilliped with spines, palp article
2 expanded, article 3 elongate, dactyl clavate,
multisetose.
Coxae 1-2 small, somewhat similar, subrectan-
gular, well setose; coxa 3 much smaller than coxa
4, adze-shaped, posteroventrally extended; coxa
4 large, not posteroventrally extended, weakly
excavate posterodorsally; only coxae 2-5 with
simple gills; brood plates 4 pairs, slender.
Gnathopods small, dissimilar, gnathopod 1
simple, gnathopod 2 weakly chelate (parache-
late), wrists elongate, hand of gnathopod 2 weakly
mitelliform. Dactyls of pereopods 3-7 distinguish-
able, those of pereopods 3-5 large, of pereopods
6-7 vestigial; dactyl of pereopod 5 large and
spinose. Article 2 of pereopods 5-7 expanded;
pereopod 5 of haustorius form; distal articles of
pereopods 6-7 not underslung, only moderately
widened, pereopods 6-7 otherwise similar, weakly
dominating pereopod 5.
Peduncles of pleopods short, wider than long,
pleopod 2 inferior, inner rami moderately shorter
than outer. Epimeron 1 scarcely differentiated,
epimeron 2 dominantly setose. Urosomite 1
scarcely produced and poorly setose ventrally.
Rami of uropods 1-2 linguiform, inner rami re-
duced, setose marginally; peduncle of uropod 1
strongly setose laterally, of uropod 2 naked or
weakly setose laterally. Uropod 3 of ordinary
form, outer ramus dominant and biarticulate,
article 2 with 2 large apical setae. Telson short,
broader than long, entire.
DESCRIPTION.?Eyes indistinct, ocular glands
visible. Dorsolateral surface of article 1 on an-
tenna 1 furnished with longitudinal row of ar-
maments; article 2 densely setose dorsolaterally,
groups distinguishable; article 3 poorly setose.
Article 3 of antenna 2 short, poorly setose, flagel-
lum much longer than article 4 of peduncle.
Upper lip especially broad.
Right lacinia mobilis linguiform, left broader
and bifid; article 3 of mandibular palp shorter
than article 2. Lower lip with 1 cone on each
outer lobe. Inner plate of maxilla 1 with 1 short
apical and 1 large apicomedial setae; spines on
outer plate 11, mostly bifid. Inner plate of max-
illiped with 2 stout, apical spines. Coxae 1-2 well
setose. Gill 2 dominant, gill 5 about as large as
anterior lobe of coxa 5. Gnathopods 1-2 bearing
medial surficial button combs.
Pereopods 3-4 lacking secondary facial rows of
spines on article 5.
Uropods 1-2 with huge dorsal setae on pedun-
cles, rami lacking basoventral setae, medial mar-
gins of peduncles heavily setose. Telson with mid-
apical seta (unusual).
TYPE-SPECIES.?Gheegerus garbaius, new species.
COMPOSITION.?Unique.
REMARKS.?Antennal flagellar setae appear to
be apically dilated, though not to such a marked
degree as those of Narunius. Oil-immersion mi-
croscopy does not magnify the elements ade-
quately for analysis.
RELATIONSHIP.?The small coxa 3 and large
coxa 4 distinguish this genus from Urohaustorius.
The untoothed epimeron 2 with vertical row of
setae, uncleft telson and vestigial dactyls of per-
eopods 6-7 suggest an affinity with the group of
species in Urohaustorius containing U. halei, U.
metungi, and U. gunni. Gheegerus also differs from
those species in the short inner rami of uropods
1-2 and the unusual presence of a midapical
dominant seta on the telson. The broad upper lip
of Gheegerus occurs in some species of Urohaustorius,
especially in large specimens.
Gheegerus differs from Bumeralius in the presence
of dactylar spines on pereopod 5 and the absence
of basoventral setae on the outer rami of uropods
1-2.
A close affinity to Narunius is seen in the con-
formation of coxae 3-4 but Narunius bears special
paddle-shaped antennal setae and narrowed ar-
ticle 2 of pereopods 6-7.
Tuldarus and Tottungus lack dactylar spines on
pereopod 5; have coxa 3 dominant; and all setae
on the palp of maxilla 1 are fully apical.
The diagnosis of the type-species follows that
in Urohaustorius for comparative value.
108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Gheegerus garbaius, new species
FIGURES 38, 39
DIAGNOSIS.?Dorsolateral armament rows on
articles 1 and 2 of antenna 1 and article 4 of
antenna 2 composed of thin setae. Coxa 1 not
larger than coxa 2, both bearing medium setae.
Generic attributes in coxae 3 and 4. Spines on
dactyl of pereopod 5 set singly in tandem. Dactyls
of pereopods 6 and 7 vestigial. Epimeron 2
rounded, with several facial setae in vertical row.
Tooth of epimeron 3 of medium size. Inner rami
of uropods 1 and 2 much shorter than outer rami.
Dorsolateral margin of uropod 2 peduncle naked
or with thin setae. Apicolateral corner on pedun-
cle of uropod 3 with thick spines. Telson entire,
very broad, bearing dominant midapical seta.
DESCRIPTION OF FEMALE.?Holotype female
"w," 2.42 mm (Victoria): Head about 75% as
long as wide, rostrum about 10% as long as
remainder of head. Primary flagellum of antenna
1 with 6 articles, formula of long aesthetascs =
0-1-1-1-1-0, accessory flagellum 5-articulate,
thin, shortened. Facial formula of elongate setae
on article 1 of antenna 1 = 5E, 3P; of shorter
elements on article 4 of antenna 2 =
PE-PE-2E-4E (first 2 sets ventrad); on article 5
= 10E; article 3 with 3 facial setae; flagellum
with 6 articles. Right and left incisors weakly
bifid, choppers on molars with 4 cusps; formula
on right palp article 3 = 10-1-3-1, on left =
9-1-3-1. Inner plate of maxilla 1 with 1 medium
apical seta, 1 large apicomedial seta; outer plate
with 11 spines. Inner plate of maxilla 2 with 4
medial setae in submarginal row, outer plate with
1 stout and 1 small apicolateral setae. Inner plate
of maxilliped with 4 medial setae, outer plate
with 1 apicolateral seta; palp article 2 with 5
setae in facial row, article 3 with 2 setae in facial
row, dactyl with 3 long setae, apical, apicolateral,
and apicomedial, inner margin naked or with 1
setule.
Coxa 1 softly rounded apically, with 10 me-
dium to short setae; coxa 2 rectangular, with 8
setae; coxa 3 with 5 long posteroventral setae, 3
shorter posterior setae and 7 medium to short
ventral setae anterior to elongate group; coxa 4
with 8 medium ventral setae, 8-9 mixed posterior
setae.
Setal and spine formulas on pereopod 3 = 4, 6,
2-1-1-1, 5 + 3 + 1 ; pereopod 4 = 4, 6, 2-1-1, 5
+ 3 + 1 , margins lacking serrations. Article 2 of
pereopods 5-7 moderately setose posteriorly, ar-
ticle 2 of pereopod 5 lacking mediofacial brush,
dactyl with 4-5 spines in tandem.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2 and 0, 2 and 1; segmental formulas =
16-10, 12-9, 14-13 (inner proliferate); basal setal
formulas = 12-2-2-3, 7-1-1-1, 9-2-2-3.
Epimeron 2 with 4 setae in vertical posterior
row.
Lateral margin on peduncle of uropod 1 with
3 setae, apicolateral corner with I setule, of uro-
pod 2 with 1 long seta and 1 setule both apico-
lateral, dorsal setae = 1 and 1, setae of rami = 6-
2 and 6-2, inner rami extending 46%) and 56%
along outer. Apicolateral corner of peduncle on
uropod 3 with 2 spines, medial margin naked,
ventromedial apical row with 6 mixed setae; lat-
eral marginal formula of article 1 on outer ramus
= Ee.
Telson about 1.33 times as wide as long, un-
cleft, with midapical seta (unusual), lateral for-
mula = P, P, P.
Gland clusters not apparent in urosome or
pereopods 5-7.
Female "g," 2.28 mm: Article 1 of antenna 1
with 5 long setae and 1 penicillate; primary fla-
gellum with 6 articles, anesthetasc formula = 0-
1-2-1-1-0. Formula on article 4 of antenna 2 =
p_p_PE-PE-4E; article 5 = 4E, 5E (and 5 in
apical vertical row); flagellum with 6 articles.
Peduncle of uropod 1 with 3 lateral setae; of
uropod 2 with 1 thin lateral seta and 1 apicolat-
eral seta; setae of rami = 6-2 and 6-2. Telsonic
formula = 1 midapical seta and 1, 1, P, P, P, or
1, P, P, P, (illus.).
Female "p," 2.12 mm: Formula on article 4
of antenna 2 = PE-PE-PE-4E. Article 1 on outer
ramus of uropod 3 with additional lateral seta.
Telsonic formula = apical seta and 1, P, P, P.
DESCRIPTION OF JUVENILE (juvenile "z," 2.11
NUMBER 360 109
KP3
W
FIGURE 38.?Gheegerus garbaius, new species, holotype, female "w," 2.42 mm (g ? female "g,"
2.28 mm; p = female "p ," 2.12 mm; z ? juvenile "z," 2.11 m).
110 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 39.?Gheegerus garbaius, new species, holotype, female "w," 2.42 mm (p = female "p ,"
2.12 mm; Z = gill).
NUMBER 360 111
mm).?Formula on article I of antenna 1 = 3E
and 2P; primary flagellum with 6 articles, aes-
thetass like female "g"; accessory flagellum 4-
articulate. Formula on article 4 of antenna 2 =
PE-PE-P-4E, on article 5 = 9; flagellum 6-artic-
ulate. Epimeron 2 with 3 setae. Peduncle of uro-
pod 1 with 2 lateral setae, of uropod 2 with 1
small, apicolateral seta; dorsal setae = 1 and 1;
setae of rami = 6-2 and 5-1. Telsonic formula
= 1 apical seta and P, P, P.
ILLUSTRATIONS.?Mouthparts generally like
Urohaustorius pulcus, except where noted.
HOLOTYPE.?NMV, female "w," 2.42 mm (il-
lus.).
TYPE-LOCALITY.?QM W.6385, station 15
QUS, Middle Banks, Moreton Bay, Queensland,
Sep 1972, 6.7 m, sand.
VOUCHER MATERIAL.?Type-locality: female
"g," 2.28 mm (illus.); female "p," 2.12 mm (il-
lus.); juvenile "z," 2.11 mm (illus.) and 3 other
specimens.
MATERIAL.? 1 sample (7).
DISTRIBUTION.?Queensland, Moreton Bay, 6.7
m, sand.
Nanmius, new genus
DIAGNOSIS.?Rostrum weak. Peduncle of an-
tenna 1 short, stout, articles 2 and 3 of peduncle
progressively shortened, scarcely geniculate, both
flagella elongate. Antenna 2 of haustorius form,
article 4 expanded, article 5 small, articles 4-5
with facial armaments, article 4 with long ventral
setae and subventral clusters of bifid setae. Aes-
thetascs simple, flagellar setae paddle-shaped.
Mandibular incisors overextended, thin, essen-
tially untoothed; right and left laciniae mobiles
not alike; rakers serrate, 2 on right mandible, 3
on left; molar broad and bulky, not strongly
extended, with weak triturative surface, with cus-
pidate accessory chopper; setae of palp article 3
awned. Mandibular lobes of lower lip well devel-
oped. Inner plate of maxilla 1 small, slender,
poorly setose, outer plate with 11 spines, palp
slightly shortened, uniarticulate, palpar setae at-
tached in 2 sets. Inner plate of maxilla 2 with
weakly submarginal oblique row of setae. Maxil-
lae and maxillipeds lacking baler lobes. Outer
plate of maxilliped with spines, palp article 2
expanded, article 3 elongate, dactyl clavate, mul-
tisetose.
Coxae 1-2 small, 3-4 larger, coxae 1-2 about
equal in size, most of coxa 2 hidden by coxa 3,
latter subequal in size to coxa 4 and hatchet
shaped, extended posteroventrally; only coxae 2-
5 with simple gills; brood plates 4 pairs, slender.
Gnathopods small, dissimilar, gnathopod 1
simple, gnathopod 2 weakly chelate (parache-
late), wrists elongate, hand of gnathopod 2 weakly
mitelliform. Dactyls of pereopods 3-7 distinguish-
able, those of pereopods 3-5 large, those of per-
eopods 6-7 small, dactyl of pereopod 5 large and
spinose. Article 2 of pereopod 5 expanded; per-
eopod 5 of haustorius form, distal articles of
pereopods 6-7 not underslung, only moderately
widened; pereopods 6-7 otherwise similar, not
dominating pereopod 5; article 2 of pereopod 6
narrow, of pereopod 7 expanded proximally, nar-
rowed distally.
Peduncles of pleopods short, wider than long,
pleopod 2 inferior, inner rami moderately shorter
than outer. Epimeron 1 scarcely differentiated,
epimeron 2 dominantly setose. Urosomite 1
scarcely produced and poorly setose ventrally.
Rami of uropods 1-2 linguiform, marginally se-
tose; peduncle of uropod 1 strongly setose lat-
erally, of uropod 2 poorly setose laterally (weak
and sparse setae). Uropod 3 of ordinary form,
outer ramus dominant and biarticulate, article 2
with 2 large apical setae. Telson short, broader
than long, weakly cleft.
DESCRIPTION.?Eyes indistinct or poorly dis-
cernible, ocular ganglia visible. Dorsolateral sur-
face of article 1 on antenna 1 furnished with
longitudinal row of armaments; article 2 densely
setose dorsolaterally, groups distinguishable, ar-
ticle 3 poorly setose. Article 3 of antenna 2 short,
poorly setose, flagellum much longer than article
4 of peduncle.
Right lacinia mobilis linguiform, left broad,
simple. Lower lip with 1 cone on each outer lobe.
Inner plate of maxilla 1 with very few short apical
112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
setae and 1 large apicomedial seta; spines on
outer plate mostly bifid or trifid. Inner plate of
maxilliped with 2 stout apical spines. Coxae 1-2
well setose. Gill 2 dominant, gill 5 smaller than
posterior lobe of coxa 5. Gnathopod 2 bearing
surficial button combs.
Pereopods 3-4 bearing secondary facial row of
spines on article 5.
Uropods 1-2 with huge dorsal setae on pedun-
cles, rami lacking basoventral setae, medial mar-
gins of peduncles heavily setose.
TYPE-SPECIES.?Narunius tallerkus, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This genus is very similar to
Urohaustorius but differs in the relatively greater
size of coxa 4 compared with coxa 3; the narrow-
ing of article 2 on pereopods 6-7; the develop-
ment of weak secondary facial spination on article
5 of pereopods 3-4; and in the distinctive paddle-
shaped flagellar setae, occuring on both antennae
1-2. Like Urohaustorius perkeus, coxae 1-2 are well
setose but otherwise U. perkeus and N. tallerkus
have many distinctions.
For the sake of comparison to species in Uro-
haustorius, the diagnosis of N. tallerkus follows the
formula for Urohaustorius.
Narunius tallerkus, new species
FIGURES 40-43
DIAGNOSIS.?Telson distinctly cleft, emargina-
tion weak and gaping. Apicolateral corner of
peduncle on uropod 3 with thin setae. Inner rami
of uropods 1-2 much shorter than outer rami.
Dorsolateral margin of peduncle on uropod 2
with small setae. Spines on dactyl of pereopod 5
set singly in tandem. Dactyls of pereopods 6-7
weak. Dorsolateral armament rows on articles 1-
2 of antenna 1 and article 4 of antenna 2 com-
posed of thin setae. Epimeron 2 rounded, with
7+ facial setae in vertical row. Tooth of epimeron
3 obsolescent. Coxa 1 not larger than coxa 2;
coxae 1-2 bearing only tiny setules; otherwise
with generic attributes in paddle setae, coxa 3,
pereopods 3-4 and 6-7.
DESCRIPTION OF FEMALE.?Holotype female
"p," 3.36 mm (New South Wales): Head about
65% as long as wide, rostrum mabout 10% as long
as remainder of head. Primary flagellum of an-
tenna 1 with 9-10 slightly thickened articles, each
ringed with numerous deeply curved thin setae,
each paddle-shaped at apex, plus normal aesthe-
tasc formula = 0-0-2-2-2-2-2-1-1-0; flagellum
9-articulate. Facial formula of setae on article 1
of antenna 1 = 6 (mixed setae); on article 4 of
antenna 2 = 3-7-5-26; on article 5 = 17; article
3 with 10 facial setae; flagellum with 11 articles
also furnished with paddle setae. Right and left
incisors apically simple, with greatly subapical
boss, choppers on molar with 3 cusps; formula on
right palp article 3 = 16-1-3-1, on left = 15-1-
3-1. Inner plate of maxilla 1 with 1 short apical
seta, 1 large apicomedial seta; outer plate with
11 spines. Plates of maxilla 2 more slender than
in Urohaustorius. Inner plate of maxilliped with 6
medial setae, apical spines shorter than in Uro-
haustorius, outer plate with 1 apicolateral seta,
palp article 2 with 7 setae in facial row, article 3
with 4 setae in facial row and 5 ventrolateral row;
palp article 4 as in U. pulcus.
Coxae 1-2 each in form of parallelogram,
strongly setose ventrally; coxa 3 with most of
ventral margin from anterior edge to M. 90,
furnished with numerous short setae (unlike Uro-
haustorius), then 5 long ventral setae, posterior
margin with 10 setae; coxa 4 with 10 long ventral
setae and 11 posterior setae. Setae of brood plates
vestigial.
Wrists of gnathopods anteriorly hirsute.
Setal and spine formulas on pereopod 3 = 9, 8,
2-1-2-2-1-1, or 2-1-1-2-1-1, 6 + 4 + 1, or 5
+ 4 + 1 (note 1 of 4 = seta); on pereopod 4 = 8
and 2 anterior, 7, 2-1-2-2-1, or 2-1-2-1-1-1, 5
+ 4 + 1 , posterior margin of articles 5-6 with
partially doubled row of mostly large even serra-
tions (like U. pulcus). Article 2 of pereopods 5-7
densely setose posteriorly; article 2 of pereopod 5
with mediofacial duplicate row of setae even
denser than on posterior margin, anterior margin
densely prickled, dactyl with 7 spines in tandem.
Peduncular spine formulas of pleopods 1-3 =
NUMBER 360 113
Gl
FIGURE 40.?Narunius tallerkus, new species, holotype, female "p," 3.36 mm (k = male "k," 2.49
mm; n = female "n," 2.73 mm; Y = brood plate; Z = aesthetasc).
114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 41.?Narunius tallerkus, new species, holotype, female "p," 3.36 mm (c = juvenile "c,'
1.48 mm; k = male "k," 2.49 mm; u = female "u," 3.84 mm; Y = pleopod).
NUMBER 360 115
P6
FIGURE 42.?Narunius tallerkus, new species, holotype, female "p , " 3.36 mm (Z = brood plate).
2 and 1, 2 and 0, 2 and 1; segmental formulas =
22-14, 17-11, 20-17 (proliferate); basal setal for-
mulas = 14-3-4-4, 5-1-1-1, 11-2-5-4.
Epimeron 2 with 7 large and 1 small setae in
vertical facial row.
Lateral margin of peduncle on uropod 1 with
6 setae and weakly extraordinary gap between
ultimate and penultimate setae, apicolateral cor-
ner with 1 setule; of uropod 2 with 3 short wiry
setae, apicolateral corner with 2 similar setae;
dorsal setae = 3 and 1; setae of rami = 9-6 and
(9-10)-5, inner rami extending 70% and 65%
along outer. Apicolateral corner of peduncle on
uropod 3 with 2 setules, medial margin with 4
setules in tandem, ventromedial apical rows with
about 21 mixed setae; setal formula on lateral
margin of article 1 on outer ramus (right side
with additional long seta as shown by dots in
illustration); formula = e-e-e-E-E-E-E-Ee.
Telson about 1.2 times as wide as long, cleft
about 0.14 of its length, setal formula = 2P, 2, P,
2.
Lateral gland clusters developed only very
weakly in pleonite 4.
Female "y," 2.79 mm (Victoria): Formulas of
armament rows on article 1 of antenna 1 = 3 +
2 tiny distals; on article 4 of antenna 2 = 4-4-2-
5-17; on article 5 = 14+; flagellar articles on
antenna 1, primary = 9, accessory = 8, antenna
2 = 9. Left lacinia mobilis weakly bifid; formula
on right and left palp article 3 = 12-1-3-1.
Ventral margin of coxa 3 almost straight. Spine
and setal formulas on pereopod 3 = 8, 7, 2?1?2?
1-1, 5 + 4 + 1; on pereopod 4 = 9 (= 2 anterior),
7, 2-1-2-1-1, 4 + 3 + 1 . Dactyl of pereopod 5
with 6 spines. Epimeron 2 with 7 setae. Peduncle
of uropod 1 with 6 lateral setae; of uropod 2
naked (1 apical setule only); dorsal setae = 3 and
1; setae of rami = 9-6 and 8-3. Article 1 on outer
ramus of uropod 3 with 8 long lateral setae. Brood
plates like holotype.
Female "f," 2.94 mm (Botany Bay):
Epimeron 2 with 8 setae. Telsonic formula = 2P,
1, P, 2; brood plates of normal form, with elongate
setae.
Female "n," 2.73 mm (New South Wales):
Left lacinia mobilis weakly bifid (illus.). Serra-
tions on articles 5-6 of pereopods 3-4 very weak;
telsonic formula = 2P, 1, P, 1 (illus.), shape
unusual.
Female "u," 3.84 mm (Queensland): Dactyl
of pereopod 5 with 6 spines in tandem, plus 1
facial spine paired with spine 5 (illus.). Epimeron
2 with 8 large and 2 tiny setae. Peduncle of
uropod 1 with 7 lateral setae; outer ramus like
holotype; peduncle of uropod 2 with only 2 api-
kRS
FIGURE 43.?Narunius tallerkus, new species, holotype, female "p," 3.36 mm (c = juvenile female
"c," 1.48 mm; h = male "h," 3.54 mm; k = male "k," 2.49 mm; n = female "n," 2.73 mm; Y
= pleopod).
NUMBER 360 117
colateral setules. Telsonic setal formula = 2P, 3,
P, 3 or 2P, 2, P, 3.
Female "x," 3.41 mm (Queensland): Dactyl
of pereopod 5 like female "u."
DESCRIPTION OF MALE.?Male "k," 2.49 mm
(New South Wales): Formulas of armaments on
article 1 of antenna 1 = 4; on article 4 of antenna
2 = 3-4-4-14; on article 5 = 8; dorsal article 4
also with short setae as in female; flagellar for-
mulas, antenna 1 primary flagellum = 9, acces-
sory flagellum = 7, antenna 2 = 8, primary
flagellum thickened as in female. Proximal hump
on mandibular incisors obsolescent; formula on
right palp article 3 = 10-1-3-1, on left = 11 ? 1?
4-1. Mandibular lobes of lower lip blunter than
in female (female like U. pulcus illustration). Max-
illa 2 like female but with 6 setae in medial setal
row. Inner plate of maxilliped like female, includ-
ing apical spines; palp article 2 with 5 facial
setae; article 3 with 3 facial and 3 ventrolateral
setae or facial row composed of 3 basal and 1
distal members; dactyl with 6 setae. Coxae 1 with
11 setae; coxa 2 with 6 setae; coxa 3 illustrated,
more elongate than in female. Formulas for both
pereopods 3-4, posterior article 5 = 2-2-1-1,
article 6 = 4 + 3 + 1; articles 5-6 lacking
serrations on pereopod 3, weakly developed on
pereopod 4. Dactyl of pereopod 5 shorter than in
female, with 5 spines (illus.); article 2 with fewer
submarginal setae. Article 4 of pereopod 6 also
like female, with only 2 posterior setae; postero-
ventral corner of coxa 7 with only 1 long seta.
Epimeron 2 with 4 setae; epimeron 3 with only 2
ventral setules total. Peduncle of uropod 1 with
5 lateral setae, of uropod 2 with 1 midlateral
setule and 1 apical setule; dorsal setae = 2 and 1,
setae of rami = 7-4 and 7-2. Formula of lateral
small setae on article 1 of outer ramus on uropod
3 = Ee-E-E-E-Ee. Telsonic formula = 2P, 1, P,
1 (illus.).
Male "h," 3.54 mm (robust): Primary flagel-
lum of antenna 1 with 11 articles, accessory fla-
gellum with 10, flagellum of antenna 2 with 12
articles; armament formula on article 4 of an-
tenna 2 = 5-9-8-21. Right molarial chopper with
at least 7 cusps; formula on right mandibular
palp article 3 = 15-1-3-1. Coxa 3 with 8 long
ventral and 13 posterior setae. Coxa 4 with 29
plumose setae total plus simple anteroventral
seta. Spine and setal formulas on pereopod 3 =
11,9 (and 2 anterior), 6 + 5 + 1; of article 6 on
pereopod 4 = 11-14, 10, 5 + 5 + 1. Gill of
pereopod 5 about as large as posterior lobe of
coxa 5. Dactyl of pereopod 5 with 7 spines in
tandem and facial spine forming pair with spine
5. Epimeron 2 with 14 setae (possibly more, badly
occluded). Peduncle of uropod 1 with 7 lateral
setae, of uropod 2 with 4 lateral setae, setae of
rami = 9-6 and 10-6. Venter of peduncle on
uropod 3 with 21 setae, uropod 3 otherwise illus-
trated. Telsonic setal formula = 4, P, 2-3, 2P
(illus.).
Juvenile female "c," 1.48 mm (Western
Port): Armament formulas of article 1 on an-
tenna 1 = 3; on article 4 of antenna 2 = 1-1-2-
1-2-8; article 5 = 6 (highly distal); article 3 = 3;
flagellar articles, primary of antenna 1 = 7, ac-
cessory = 6, antenna 2 = 6, aesthetascs of primary
flagellum on antenna 1 = 0-1-2-2-1-1-0. Right
lacinia mobilis bifid, left weakly bifid, chopper
cusps = 3, very elongate; formula on right and
left palp article 3 = 7-1-3-1. Medial row on
inner plate of maxilla 2 with 4 setae. Palp article
2 of maxilliped with 4 facial setae, article 3 with
2 and 3 ventrolateral setae, article 4 with 5 setae
(2 long apical, 1 longsubapical, 2 short marginal).
Coxa 1 with 7 setae; coxa 2 with 4 setae; coxa 3
with 14 short ventral setae, 5 long posteroventrals,
and 3 dorsoposterior setae; coxa 4 with 11 long
setae plus 1 anteroventral setule. Spine and setal
formulas on pereopod 3 = 3, 3, 2-1-1,3 + 2 + 1;
pereopod 4 = 4, 2, 2-1-1, 3 + 2 + 1, spines with
large sharp cusps. Pereopods 5-7 illustrated (per-
eopod 6 composite of right and left). Epimeron 2
with 3 setae. Uropods 1-3 and telson illustrated,
uropod 3 with 2 short medial setae on peduncle
and 7-8 ventral setae.
HOLOTYPE.?AM, female "p," 3.36 mm (illus.).
TYPE-LOCALITY.?EBS 148, 500 yards (455 me-
ters) off Towra Point, New South Wales, 30 Nov
1972, 3 m, sand and shell grit.
VOUCHER MATERIAL.?Type-locality: male
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
"k," 2.49 mm (illus.); female "n," 2.73 mm (il-
lus.); female "a," 2.72 mm; male "v," 2.38 mm;
EBS Botany Bay 1025: female "f," 2.94 mm;
CPBS A6/5: juvenile female "c," 1.48 mm (illus.);
CPBS 26 N/2: female "y," 2.79 mm; AM
P.23293: male "h," 3.54 mm (illus.); QM W.6384:
female "u," 3.84 mm (illus.); female "x," 3.41
mm (illus.).
MATERIAL.?CPBS, 4 samples from 4 stations
(4); PPBES, 17 samples from 8 stations (172);
EBS, 2 samples from 2 stations (5); AM, 2 samples
(2); QM, 3 samples (12).
DISTRIBUTION.?Victoria, Western Port and
Port Phillip Bay to Moreton Bay, Queensland, 3-
22 m, sand, silty sand, sand and shell, sand and
weed.
Tuldarus, new genus
DIAGNOSIS.?Rostrum weak. Peduncle of an-
tenna 1 short, stout, articles 2 and 3 of peduncle
progressively shortened, weakly geniculate, pri-
mary flagellum elongate, accessory flagellum well
developed or weak. Aesthetascs simple. Antenna
2 of haustorius form, article 4 expanded, article
5 small, articles 4-5 with facial armaments, article
4 with long ventral setae and subventral clusters
of bifid setae. Mandibular incisors overextended,
thin, essentially untoothed; right and left laciniae
mobiles not alike; rakers serrate or not, 1-2 on
right mandible, 2-3 on left, 1 raker on each side
often rudimentary; molar broad and bulky, not
strongly extended, with weak triturative surface,
accessory chopper poorly developed; setae of palp
article 3 awned. Mandibular lobes of lower lip
well developed. Inner plate of maxilla 1 small,
slender, poorly setose, outer plate apparently with
10 spines; palp slightly shortened, uniarticulate,
palpar setae attached in 1 apical set. Inner plate
of maxilla 2 with weakly submarginal oblique
row of setae. Maxillae and maxillipeds lacking
baler lobes. Outer plate of maxilliped with spines,
palp article 2 expanded; article 3 elongate, dactyl
clavate, multisetose.
Coxae 1-2 very small but coxa 1 larger than
coxa 2; coxa 3 larger than or subequal in size to
coxa 4, coxa 3 hatchet-shaped and extended pos-
teroventrally; only coxae 2-5 with simple gills;
brood plates 4 pairs, slender.
Gnathopods small, dissimilar, gnathopod 1
simple, gnathopod 2 weakly chelate (parache-
late), wrists elongate, hand of gnathopod 2 weakly
mitelliform. Dactyls of pereopods 3-7 distinguish-
able, large, dactyl of pereopod 5 lacking spines
but occasionally bearing apical setae. Article 2 of
pereopods 6-7 narrow (type) to strongly ex-
panded; of pereopod 5 strongly expanded, latter
of haustorius form, distal articles of pereopods 6-
7 not underslung, only moderately widened, per-
eopods 6-7 otherwise similar, weakly dominating
pereopod 5.
Peduncles of pleopods short, wider than long,
pleopod 2 inferior, inner rami moderately shorter
than outer. Epimeron 1 scarcely differentiated,
epimeron 2 dominantly setose. Urosomite 1
scarcely produced and poorly setose ventrally.
Rami of uropods 1-2 linguiform, though inner
rami often reduced or vestigial, marginally setose;
peduncle of uropod 1 strongly setose laterally, of
uropod 2 generally lacking long setae laterally.
Uropod 3 of ordinary form, outer ramus domi-
nant and biarticulate, article 2 with 2 large apical
setae. Telson short, broader than long or not
greatly longer than broad, entire.
DESCRIPTION.?Eyes indistinct or poorly visible
or occasionally well developed, ocular ganglia
visible. Dorsolateral surface of article 1 on an-
tenna 1 furnished with longitudinal row of ar-
maments; article 2 densely setose dorsolaterally,
groups distinguishable; article 3 poorly setose.
Article 3 of antenna 2 short, poorly setose, flagel-
lum much longer than article 4 of peduncle.
Right lacinia mobilis linguiform, left broader
and often bifid. Lower lip usually with 1 cone on
each outer lobe. Inner plate of maxilla 1 with
very few short apical setae and 1 larger apico-
medial seta; spines on outer plate mostly bifid.
Inner plate of maxilliped with 2 stout apical
spines. Coxae 1-2 poorly setose. Gill 2 dominant,
gill 5 rarely larger, often much smaller than
posterior lobe of coxa 5. Gnathopod 2 bearing
surficial button combs.
NUMBER 360 119
Pereopods 3-4 lacking extensive secondary fa-
cial rows of spines on article 5.
Uropods 1-2 lacking huge dorsal setae on pe-
duncle, rami lacking basoventral setae, medial
margins of peduncles usually heavily setose.
TYPE-SPECIES.?Tuldarus cangellus, new species.
COMPOSITION.? Tuldarus barinius, new species.
RELATIONSHIP.?This genus differs from Uro-
haustonus in the absence of large stiff spines on the
dactyl of pereopod 5 and the fully apical position
of all setae on the palp of maxilla 1. These small
differences are reinforced by the tendency for the
inner ramus of uropods 1-2 to become fused to
the peduncle or lost entirely; for 1 of the mandib-
ular rakers to exist in rudimentary form; and for
the tendency towards complete domination by
spines in the armament row on article 4 of an-
tenna 2. The type-species is characterized by loss
of dominance by coxa 3 and by article 2 on
pereopods 6-7; while the second species is char-
acterized by total loss of inner rami on uropods
1-2.
The type-species of Tuldarus weakly replicates
some tendencies seen in Narunius, such as the
narrowed article 2 of pereopods 6-7 and the small
coxa 3, but Tuldarus differs from Narunius in the
absence of stout dactylar spines on pereopod 5
and of the special oared setae on the antennae.
Key to Species of Tuldarus
Inner rami of uropods 1-2 half as long as outer rami
T. cangellus, new species
Inner rami of uropods 1-2 absent T. barinius, new species
Tuldarus cangellus, new species
FIGURES 44-46 (part)
DIAGNOSIS.?Inner rami of uropods 1-2 artic-
ulate, about half as long as outer rami. Article 2
of pereopods 6-7 of narrow form (see illustra-
tions). Tooth of epimera 1-2 sharp and attenuate.
Dactyls of pereopods 3-4 with accessory tooth
vestigial. Posterior lobe of coxa 3 sharp, attenuate,
coxa 4 also with attenuate posteroventral lobe.
DESCRIPTION OF MALE.?Holotype male "i,"
4.12 mm: Head about 60% as long as wide,
rostrum about 7% as long as remainder of head.
Primary flagellum of antenna 1 with 12 articles
furnished densely with long aesthetascs in for-
mula from proximal to distal: 5-5-6-6-6-6-6-5-
5-3-2-0, accessory flagellum 7-articulate. Facial
formula of armaments on article 1 of antenna 1
= 5 setae, on article 2 = 4 spines and 11 setae, on
article 4 of antenna 2 = mixture of 14 spines, 9
setae, 2 penicillates, on article 5 = mixture of 4
spines and 6 setae (not counting transverse apical
row), article 3 with 2 facial spine-setae; flagellum
with 8 articles. Right incisor simple, left weakly
bifid; right lacinia mobilis weakly trifid, left bifid;
right rakers = 1 plus 1 rudimentary, left = 2 plus
1 rudimentary; chopper humps simple; palp ar-
ticle 2 thin, with only 2 setae, formula on right
and left article 3 = 6-1-3-1. Inner plate of max-
illa 1 with 1 apical seta, 1 large apicomedial seta;
outer plate apparently with 10 spines, several
with as many as 3 lateral cusps. Inner plate of
maxilla 2 with 5 setae in medial row. Inner plate
of maxilliped with 6 medial setae, outer plate
with 1 apicolateral seta; palp article 2 with 3
setae in facial row, article 3 with 2 setae in facial
row, dactyl with 3 long and 3 short setae.
Coxa 1 slipper-shaped, attenuate but apically
rounded; coxa 2 much shorter, broad, subrectan-
gular; coxa 3 not much larger than coxa 4, with
3 long ventral setae near posterior corner; coxa 4
with 2 long ventral setae and 9 mixed posterior
setae.
Setal and spine formulas on pereopod 3 = 5, 2,
2-1-1-1, 4 + 2 + 1 ; pereopod 4 = 5, 2, 2-1-1-1,
4 + 3 + 1, note absence of posterior setae on
article 5 of pereopod 3, posterior margin of article
5 with single row of serrations similar to U. pulcus.
Article 2 of pereopods 5-6 with sparse setal den-
sities, of pereopod 7 with few weak setules, pos-
120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
VX1
FIGURE 44.? Tuldarus cangellus, new species, holotype, male "i,
mm).
4.12 1mm (& = male "v," 3.96
NUMBER 360 121
FIGURE 45.?Tuldarus cangellus, new species, holotype, male "i ," 4.12 mm (a = posterior margin
rolled inward; v = male "v," 3.96 mm).
122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
X2 VX1
FIGURE 46.?Upper: Tuldarus cangellus, new species, holotype, male "i," 4.12 mm (v = male "v,
3 .% mm; Z = gill). Lower: Tuldarus barinius, new species, holotype, female "r," 1.78 mm (w ?
female "w," 1.62 mm; x = male "x," 1.65 mm; Y = eye).
NUMBER 360 123
teroventral lobe on pereopod 7 weak, on pereopod
6 obsolescent; dactyl of pereopod 5 with several
anterior humps.
Peduncular spine formulas of pleopods 1-3 =
2 and 1, 2, and 0, 2 and 0; segmental formulas
= 16-10, 16-9, 14-9; basal setal formulas = 9-2-
2-2,4-1-1-2,4-2-2-2.
Epimeron 2 with 4 setae in horizontal but
weakly oblique row.
Lateral margin on peduncle of uropod 1 with
4-5 setae and 1 apicolateral setule, of uropod 2
with 2 apicolateral setules only; dorsal setae ab-
sent; setae of rami = 8-5 and 7-4; inner rami
extending 67% and 60% along outer. Apicolateral
corner of peduncle on uropod 3 with 2 setules,
medial margin with 1 setule, ventromedial apical
row with 5-6 setae; formula on lateral margin of
article 1 on outer ramus = ee-Ee-Ee-Ee-Ee;
uropod 3 especially slender.
Telson about 1.3 times as wide as long, apex
convex, setal formula = 2P, 1, P, 1/2P, 1, P, 1,
distalmost P seta submarginal.
Urosomal and telsonic glands not discernible.
Male "v," 3.96 mm: Rostrum slightly sharper
and more protuberant than in holotype. Primary
flagellum of antenna 1 with 11 articles, accessory
with 6. Article 4 of antenna 2 with 23 elements in
armament row, formula = P, S, P, S, P, SS-
SESSS-P-SE-SESE-SEES; flagellum with 8 ar-
ticles. Spine and setal formulas on pereopod 3 =
5, 2, 2-1 or 2-1-1, 4 + 3 + 1 ; pereopod 4 = 5, 2
or 3, 2-1, 4 + 3 + 1. Article 5 of pereopod 6 with
combed apex as on articles 5-6 of pereopods 3-4.
Uropod 1 with 4 lateral setae; setae on rami of
uropods 1-2 = 7-5 and 6-4. Setal placement on
telson somewhat aberrant (illus.).
DESCRIPTION OF FEMALE (female "z," 3.31
mm): Brood plates small and lacking setae; ar-
ticle 1 of antenna 1 with 2 long and 1 short
plumose setae + 2P; article 2 with 2 short stout
spines, 4 long slender simple setae, and 5 long
plumose dorsolateral setae; aesthetascs short
(about twice as long as flagellar articles), formula
on primary flagellum = 0-1-2-2-2-1-1-0; acces-
sory flagellum not reduced, 5-articulate; dactyl
on pereopod 5 with 1 long subapical seta (unlike
male); epimeron 2 with 3 facial setae; lateral
marginal formula on article 1 of outer ramus on
uropod 3 = e-Ee-Ee; formula on telson = 2P, P,
1/2P, P, 1, 1.
ILLUSTRATIONS.?Pereopod 5 of holotype com-
posite of right and left sides, pereopod 5 of male
"v" thus illustrated for confirmation; posterior
and apical margins of coxa 1 curled, difficult to
flatten, coxa 1 of holotype shown with 3 posterior
edges as fully rolled, partially unrolled in situ,
and flattened drastically; coxa 1 of male "v"
shown as apically rolled with flattened extension
in hatched line.
HOLOTYPE.?NMV, male "i," 4.12 mm (illus.).
TYPE-LOCALITY.?CPBS 5IN, Western Port,
Victoria, 30 Mar 1965, 16.5 m, sand and mud.
VOUCHER MATERIAL.?Type-locality, male
"v," 3.96 mm (illus.); AM P.23326, female "z,"
3.31 mm.
RELATIONSHIP.?The second species, to follow,
is compared with this, the model of the genus.
MATERIAL.?CPBS, 4 samples from 3 stations
(4); AM 1 sample (1).
DISTRIBUTION.?Victoria, Western Port to New
South Wales, 3.5 km S of Hunter River, 11-28
m, sand, sand and mud.
Tuldarus barinius, new species
FIGURES 46 (part)-48
DIAGNOSIS.?Inner rami of uropods 1-2 absent.
Article 2 of pereopods 6-7 of broad form (see
illustrations). Tooth of epimera 1-2 blunt and
not attenuate. Dactyls of pereopods 3-4 with
sharp accessory tooth. Posterior lobe of coxa 3
blunt, not attenuate; coxa 4 lacking posteroven-
tral lobe.
DESCRIPTION OF FEMALE.?Holotype female
"r," 1.78 mm: Head about 70% as long as wide,
rostrum about 10% as long as remainder of head.
Primary flagellum of antenna 1 with 6 articles
furnished sparsely with long aesthetascs in for-
mula: 0-0-1-1-1-0; accessory flagellum 3-artic-
ulate. Facial formula of armaments on article 1
of antenna 1 = 4 setae; on article 2 = 8 setae or
124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 47.?Tuldarus barinius, new species, holotype, female "r," 1.78 mm (b = female "b,
1.57 mm; x ? male "x," 1.65 mm; Z = gill).
NUMBER 360 125
FIGURE 48.?Tuldarus bannius, new species, holotype, female "r," 1.78 mm (u> ? female "w,"
1.62 mm; x = male "x," 1.65 mm).
spine-setae, no distinctly thick spines; on article
4 of antenna 2 = mixture of 10 spines, 1 seta and
2 penicillates in 2 groups; on article 5 = 6 spine-
setae; article 3 with 1 seta; flagellum with 6
articles. Right incisor simple, left bifid; right
lacinia mobilis bifid, left weakly trifid; right rak-
ers = 1, left = 2; chopper humps simple; palp
article 2 stout with 1 seta, formula on left palp
article 3 = 5-1-2-1 (right broken). Inner plate of
maxilla 1 with 1 apical seta, 1 large and closely
contiguous apicomedial seta; outer plate appar-
ently with 10 spines. Inner plate of maxilla 2 with
2 setae in medial row. Inner plate of maxilliped
with 4 medial setae; outer plate lacking apicolat-
eral seta (illus.); palp article 2 with 2 setae in
facial row set near apex, article 3 lacking facial
setae, dactyl with only 1 of short marginal seta-
group.
Coxa 1 slipper-shaped, apically rounded but
not attenuate; coxa 2 much shorter, weakly ovate;
coxa 3 much larger than coxa 4, with only 1 long
seta situated at blunt posteroventral corner, ven-
tral margin lined sparsely with setules; coxa 4
with 4 ventral setules, naked posteriorly; brood
plates slender.
Setal and spine formulas on pereopod 3 = 3, 0,
2-1, 3 + 3 + 1 ; pereopod 4 = 4, 3, 2-1, 3 + 3 +
1, posterior margin of article 5 with densely
packed serrations. Article 2 of pereopods 5-7
almost evenly but moderately setose, setae on
pereopod 7 mostly short, posteroventral lobe on
126 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
pereopods 6-7 broad but very shallow; dactyl of
pereopod 5 very finely serrate anteriorly.
Peduncular spine formulas on pleopods 1-3 =
2 and 0, 2 and 0, 2 and 0; segmental formulas =
9-6, 9-6, 9-5; basal formulas = 6-2-1-2, 3-1-1-
1,4-1-1-2.
Epimeron 2 with 1 facial seta midposteriorly;
epimeron 3 with 1 facial seta in middle of plate.
Lateral margin on peduncle of uropod 1 with
3 setae, extraordinary gap between ultimate and
penultimate, apicolateral setule set far ventrally;
of uropod 2 naked laterally, dorsal setae absent,
setae of rami = 5-0 and 4-0, inner rami absent.
Apicolateral corner of peduncle on uropod 3 with
2 thin spines, apicomedial corner with 1 long seta,
1 short spine together forming medial and ventral
armaments; spine formula on lateral margin of
article 1 of outer ramus = e-Ee; uropod 3 espe-
cially slender.
Telson nearly 1.2 times as wide as long, apex
weakly convex, setal formula = 2P, P, 1, 1, dis-
talmost P seta submarginal.
Glands dispersed in pleonites 1-6, telson and
uropod 3.
Female "w," 1.62 mm: Posterior margin of
coxa 4 straight. Basalmost medial peduncular
seta on uropod 1 absent. Ramus of uropod 2 with
3 setae. Telsonic setal formula = 2P, P, 1 (illus.).
DESCRIPTION OF MALE (male "x", 1.65 mm).?
Formula of aesthetascs on primary flagellum of
antenna 1 = 0-1-2-3-1-1-0. Facial formula of
armaments on article 4 of antenna 2 = 10 spines,
3 setae, and 4 penicillates; on article 5 = 3 spines
and 2 setae. Formula on mandibular palp article
3 = 5-1-2-1. Spine and setal formulas on pereo-
pod 3 = 3, 0, 2-1, 3 + 3 + 1 , pereopod 4 = 4, 3,
2-1,3 + 3 + 1. Ramus of uropod 2 with 3 setae.
Telsonic setal formula = 2P, P, 1-2 (illus.).
OBSERVATIONS.?Ventral margin of upper lip
naked. Eye capsules well developed in both sexes,
in female with tiny ommatidium in each capsule,
in male moderately packed with at least 2 pairs
of ommatidia plus other tissue possibly embryonic
ommatidia or other receptor cells, eyes reflecting
violet light at far distance, on closer view om-
matidia appearing pink.
HOLOTYPE.?NMV, female "r," 1.78 mm (il-
lus.).
TYPE-LOCALITY? CPBS 25S/5, Western Port,
Victoria, 25 Feb 1965, 9 m, muddy sand.
VOUCHER MATERIAL.?CPBS 31E/1: female
"b," 1.57 mm (illus.); female "w," 1.62 mm (il-
lus.); male "x," 1.65 mm (illus.).
RELATIONSHIP.?This species differs from the
type-species, T. cangellus, in the absence of inner
rami on uropods 1-2, the smaller and blunter
teeth of epimera 2-3, the presence of a midfacial
seta on epimeron 3, the blunt posteroventral lobe
of coxa 3, and the absence of a posteroventral
lobe on coxa 4, the broader article 2 on pereopods
6-7, the stronger accessory tooth on the dactyls
of pereopods 3-4, the denser anterior serrations
on the dactyl of pereopod 5, the absence of facial
setae on article 5 of pereopod 3, and the less
attenuate coxa 1.
MATERIAL.?CPBS, 2 samples from 2 stations
(5).
DISTRIBUTION.?Victoria, Western Port, 9-12
m, muddy sand.
Tottungus, new genus
DIAGNOSIS.?Rostrum weak, but head large
and cowl-like. Peduncle of antenna 1 short, stout,
articles 2 and 3 progressively shortened, weakly
geniculate; primary flagellum elongate, accessory
flagellum weak. Antenna 2 of haustorius form,
article 4 expanded, article 5 small, articles 4-5
with facial armaments, article 4 with long ventral
setae and subventral clusters of bifid setae. Aes-
thetascs simple. Mandibular incisors overex-
tended, thin, essentially untoothed; right and left
laciniae mobiles not alike; rakers usually serrate,
2 on right mandible, 3 on left, 1 raker usually
rudimentary; molar broad and bulky, not
strongly extended, with weak triturative surface,
with simple accessory chopper; setae of palp ar-
ticle 3 awned. Mandibular lobes of lower lip well
developed. Inner plate of maxilla 1 small, slender,
poorly setose, outer plate with 11 spines, palp
slightly shortened, uniarticulate, palpar setae at-
tached in 2 sets. Inner plate of maxilla 2 with
NUMBER 360 127
weakly submarginal oblique row of setae. Maxil-
lae lacking baler lobes, maxillipeds with weak
baler lobe basally. Outer plate of maxilliped with
spines; palp article 2 expanded, article 3 elongate,
dactyl clavate, multisetose.
Coxae 1-2 small, coxa 1 larger than coxa 2;
most of coxa 2 hidden by coxa 3; latter larger
than coxa 4, adze-shaped, and extended postero-
ventrally; only coxae 2-5 with simple gills; brood
plates 4 pairs, slender.
Gnathopods small, dissimilar, gnathopod 1
simple, gnathopod 2 subchelate, wrists elongate,
hand of gnathopod 2 weakly mitelliform. Dactyls
of pereopods 3-7 distinguishable, large; dactyls of
pereopods 5-7 in female with 1 or more elongate,
subapical setae. Pereopod 5 of haustorius form,
article 2 expanded; article 2 of pereopods 6-7 of
only moderate width, distal articles not under-
slung, only moderately widened; pereopods 6-7
otherwise similar, scarcely dominating pereopod
5.
Peduncles of pleopods short, wider than long,
pleopod 3 inferior, inner rami moderately shorter
than outer. Epimeron 1 scarcely differentiated,
epimeron 2 dominantly setose. Urosomite 1
scarcely produced and poorly setose ventrally.
Outer rami of uropods 1-2 linguiform, marginally
setose, inner rami short, fused to peduncle; pe-
duncle of uropod 1 strongly setose laterally, of
uropod 2 with only 2 apicolateral setule. Uropod
3 of ordinary form, outer ramus dominant and
biarticulate, article 2 with 3 large apical setae.
Telson short, broader than long, entire.
DESCRIPTION.?Eyes indistinct or poorly dis-
cernible, ocular ganglia visible. Dorsolateral sur-
face of article 1 on antenna 1 furnished with
organized longitudinal row of armaments; article
2 with spines and plumose setae dorsolaterally;
article 3 poorly setose. Article 3 of antenna 2
short, poorly setose, flagellum much longer than
article 4 of peduncle.
Right lacinia mobilis linguiform, left broader.
Lower lip with 1 cone on each outer lobe. Inner
plate of maxilla 1 with very short apical setae and
1 large apicomedial seta; spines on outer plate
mostly bifid. Inner plate of maxilliped with 2
stout apical spines. Coxae 1-2 poorly setose. Gill
2 dominant, gill 5 smaller than posterior lobe of
coxa 5. Gnathopod 2 bearing surficial button
combs.
Pereopods 3-4 lacking extensive secondary fa-
cial rows of spines on article 5.
Uropods 1-2 lacking huge dorsal setae on pe-
duncles, rami lacking basoventral setae, medial
margins of peduncles not heavily setose.
TYPE-SPECIES.? Tottungus tungus, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?In evolutionary sequence this
genus follows Tuldarus but differs in the develop-
ment of 3 (instead of 2) apical setae on the outer
ramus of uropod 3 and in the loss of parachelate-
ness on gnathopod 2, the palm simply being
subchelate. Other characters of unusual degree in
this genus are the cowl-like head, the extra set of
facial setae on article 5 of pereopods 3-4, and the
extreme reduction in size of setae on coxae 3-4.
Tottungus tungus, new species
FIGURES 49-51
DIAGNOSIS.?With the characters of the genus;
unique features being presence of 3 apical setae
on outer ramus of uropod 3, loss of chelateness on
gnathopod 2.
DESCRIPTION OF FEMALE.?Holotype female
"d," 2.54 mm: Head about 85% as long as wide,
rostrum about 10% as long as remainder of head.
Primary flagellum of antenna 1 with 7 articles,
formula of medium length, aesthetascs = 0-1-1-
2-1-1-0; accessory flagellum 5-articulate. Facial
formula of mixed spines and setae on article 1 of
antenna 1 = 1 plumose seta, 2 spines, 2 penicil-
lates; on article 4 of antenna 2 in 2 weakly divided
sections of 10 each, spines dominant; on article 5
= 11 (omitting transverse apical set); article 3
with 1 seta; flagellum with 8 articles. Right and
left incisors almost simple, formula on right and
left palp article 3 = 4-1-2-1. Inner plate of
maxilla 1 with 1 short apical seta, 1 large apico-
medial seta tightly contiguous; outer plate with
11 spines, occasional spine with more than 2
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 49.?Totlungus tungus, new species, holotype, female "d," 2.54 mm (a = angular edge
rolled inwards; k = female "k," 2.60 mm; p = male "p," 2.60 mm; K= baler lobe).
NUMBER 360 129
P7 ZP5
FIGURE 50.? Tottungus tungus, new species, holotype, female "d," 2.54 mm ( Z = gill).
E2
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 51.?Tottungus tungus, new species, holotype, female "d," 2.54 mm (k = female "k,
2.60 mm; p = male "p ," 2.60 mm).
NUMBER 360 131
cusps. Medial row on inner plate of maxilla 2
with 3 setae. Inner plate of maxilliped with 4
medial setae; outer plate with 1 apicolateral seta;
palp article 2 with 3 setae in facial row, article 3
with 1 facial seta; basal articles of maxilliped
broadened and with weakly developed baler lobe
bearing 1 seta.
Coxa 1 slipper-shaped, apically blunt, longer
than coxa 2; latter broad, short; coxa 3 with long
seta at posteroventral corner, ventral margin
lined with small setules; coxa 4 with posteroven-
tral bevel, ventral margin with several small se-
tules, posterior margin naked.
Palm of gnathopod 2 with side by side double
set of serrations.
Setal and spine formulas on pereopod 3 = 3, 3,
2-1, 4 + 4 + 1 ; pereopod 4 = 3, 4, 2-1, 4 + 4 +
1, posterior margin of article 5 with row of even
serrations. Article 2 of pereopods 5-7 progres-
sively with weak to weaker setal sizes posteriorly;
article 2 of pereopod 5 with weak mediofacial
brush of setae, dactyl with weak anterior serra-
tions and 2 long subapical setae; dactyl of per-
eopod 6 with 4 subapical setae, of pereopod 7
with 1 subapical seta.
Peduncular spine formulas of pleopods 1-3 =
2 and 0, 2 and 0, 2 and 0; segmental formulas =
L3-9, 12-8, 10-8; basal setal formulas = 7-1-2-
2,3-1-1-1, 1-1-1-1.
Epimeron 2 with small blunt posteroventral
tooth and horizontal row of 5 setae, anteriormost
seta small, or opposite side with only 4 large setae;
epimeron 3 with quadrate posteroventral corner.
Lateral margin on peduncle of uropod 1 with
4-5 setae, 4 small apicoventral setae (perhaps
homologous to apicolateral seta of other taxa), of
uropod 2 only with 4 apicoventral setules, no
dorsal setae; setae of outer rami = 7 and 7, inner
rami represented by small peduncular projections
bearing several long setae. Apicolateral corner of
peduncle on uropod 3 with 4 setae, medial margin
naked, apicomedial corner with 1 setule, ventro-
medial apical row with 5-6 mixed setae; formula
on lateral margin of article 1 on outer ramus =
Ee-Ee-Ee. Telson about 1.5 times as wide as
long, setal formula = 2P, P, 1, 1 plumose/2P, P,
1, 1 plumose.
Gland clusters indiscernible.
Female "k," 2.60 mm: Article 2 of antenna 1
with 7 long setae, no short spines, primary flagel-
lum with 7 articles, accessory with 5. Spine and
setal formulas of pereopod 3 = 2, 3, 2-1-1, 5 + 4
+ 1; pereopod 4 = 3, 4, 2-1-1, 5 + 4 + 1 . Outer
ramus of uropod 2 with 6 setae. Telson illustrated.
DESCRIPTION OF MALE (male "p," 2.60 mm).?
Primary flagellum of antenna 1 swollen, 11-12
articulate, formula of elongate aesthetascs =
4-4-4-5-5-5-5-5-2-1-0 or 3-2-3-3-3-6-5-5-
4-2-1-0 (some omitted in illustration); setae on
article 2 of peduncle smaller than in female. Left
lacinia mobilis 4-toothed; formula on right palp
article 3 of mandible = 4-1-2-1, left = 5-1-2-1.
Spines on outer plate of maxilla 1 with only 2
branches. Spine and setal formulas on pereopod
3 = 3, 3, 2-1-1, 4 + 4 + 1 ; pereopod 4 = 4, 4,
2-1-1-1, 5 + 4 + 1. Dactyls of pereopods 5-7
lacking setae. Epimeron 2 with 5 facial setae.
Medial peduncular processes (remnants of inner
rami) on uropods 1-2 much larger than in female
(illus.); outer ramus of uropod 2 with 7 setae.
Outer ramus of uropod 3 with 4 apical setae
(illus.); peduncle with 3 apicolateral setae. Telson
weakly concave posteriorly, setal formula = 2P,
0, 3, plumose/2P, P, 3 plumose (illus.). Urosomal
glands moderately developed in segments 1-3,
absent or indiscernible in peduncle or uropod 3
and telson. Pereopodal glands well developed in
articles 2, 5, 6 of pereopods 5-7 (some of these
not discerned in female; see female illustrations).
JUVENILES.?Telson with broadly rounded
apex, seta formula = 2P, P. Article 2 of outer
ramus on uropod 3 with only 2 apical setae.
Juveniles in brood pouch with article 2 on outer
ramus of uropod 3 almost as long as article 1.
OBSERVATIONS.?Head much more cowllike
than in other taxa of Urohaustorius group (see
illustrations); article 5 of pereopods 3-4 with
extra facial set of setae not occurring in other
taxa of Urohaustorius group (see illustration).
Well-developed eyes seen in several males (il-
lus.). Telson of several males such as male "h"
132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
weakly cleft, joined in numerous transitional
specimens to males with uncleft telson.
Dactylar setation in females variable, occasion-
ally 1 or 2 of peropods 5-7 lacking setae in certain
individuals, but at least 1 of these pereopods
always with dactylar setae. Article 2 on outer
ramus of uropod 3 occasionally with 4 setae;
occasionally third seta and always fourth seta
appearing weakly lateral rather than apical. In-
ner rami of uropods 1-2 longer in juveniles than
in adults. Most common formula of facial setae
on article 1 of antenna 1 = long plumose seta,
shorter seta, spine, 2 penicillate setules; also fre-
quently occurring = long plumose seta, 2 spines,
2 penicillate setules (= holotype, illus.). Third
seta on palp of maxilla 1 usually weakly disjunct
but 1 female from CPBS 12S/2 with all setae
flush as in Tuldarus.
HOLOTYPE.?NMV, female "d," 2.54 mm (il-
lus.).
TYPE-LOCALITY.?CPBS 12S/2, Western Port,
Victoria, 18 Mar 1965, 2.1 m, muddy sand and
weed.
VOUCHER MATERIAL.?Type-locality: female
"k," 2.60 mm (illus.); female "m," 2.46 mm; male
"p," 2.60 mm (illus.); male "r," 2.40 mm; male
"q," 2.41 mm; CPBS 25S/3: juvenile " j , " 1.2
mm; CPBS 26N/S: male "h," 2.45 mm.
MATERIAL.?CPBS 27 samples from 16 stations
(199); WPBES 14 samples from 10 stations (35).
DISTRIBUTION.?Victoria, Western Port, 2-23
m, fine to coarse sand, muddy sand, sand and
shell, sand and weed.
Dirimus, new genus
DIAGNOSIS.?Rostrum weak but head large and
cowl-like. Peduncle of antenna 1 short, stout,
articles 2 and 3 of peduncle progressively
shortened (but not much), weakly geniculate;
primary flagellum elongate, accessory flagellum
weak. Antenna 2 of haustorius form, article 4
expanded, article 5 small, articles 4-5 with facial
armaments, article 4 with long ventral setae and
subventral clusters of bifid setae. Aesthetascs sim-
ple. Mandibular incisors overextended, thin, es-
sentially untoothed; right and left laciniae mo-
biles not alike; rakers serrate, 1 on right mandible,
2 on left; molar broad and bulky, not strongly
extended, with weak triturative surface, and sim-
ple accessory chopper; setae of palp article 3
awned. Mandibular lobes of lower lip well devel-
oped. Inner plate of maxilla 1 small, slender,
poorly setose, outer plate with 9 spines, palp
shortened, uniarticulate, palpar setae attached in
1 set. Inner plate of maxilla 2 with weakly sub-
marginal oblique row of setae. Maxillae lacking
baler lobes, maxilliped with weak baler lobe bas-
ally. Outer plate of maxilliped with spines; palp
article 2 expanded, article 3 elongate, dactyl cla-
vate, multisetose.
Coxae 1-2 small, of similar size, most of coxa
2 hidden by coxa 3; latter larger than coxa 4,
adze-shaped, and extended posteroventrally; only
coxae 2-5 with simple gills; brood plates 4 pairs,
slender.
Gnathopods small, dissimilar, gnathopod 1
simple, gnathopod 2 parachelate, wrists elongate,
hand of gnathopod 2 weakly mitelliform. Dactyls
of pereopods 3-7 distinguishable, large, dactyls of
pereopods 5-7 with 1 or more elongate subapical
setae. Article 2 of pereopods 5-7 expanded, per-
eopod 5 of haustorius form; distal articles of
pereopods 6-7 not underslung, only moderately
widened, pereopods 6-7 otherwise similar,
scarcely dominating pereopod 5.
Peduncles of pleopods wider than long, pleo-
pod 3 inferior (but scarcely so), inner rami mod-
erately shorter than outer. Epimeron 1 scarcely
differentiated; epimeron 2 not facially setose and
scarcely dominating epimeron 3. Urosomite 1
scarcely produced and not setose ventrally. Outer
rami of uropods 1-2 linguiform, marginally se-
tose, inner rami short and almost fused to pedun-
cles; peduncle of uropod 1 with 2 large setae
laterally, of uropod 2 with only 1 apicolateral
setule. Uropod 3 of ordinary form, outer ramus
dominant and biarticulate, article 2 with 2 large
apical setae. Telson short, very broad, entire.
DESCRIPTION.?Eyes distinct. Dorsolateral sur-
face of article 1 on antenna 1 with weak arma-
ment row; article 2 densely setose dorsolaterally,
NUMBER 360 133
groups weakly distinguishable; article 3 poorly
setose. Article 3 of antenna 2 short, poorly setose;
flagellum much longer than article 4 of peduncle.
Right lacinia mobilis linguiform, slender; left
thin and spike-like. Lower lip with 1 cone on each
outer lobe. Inner plate of maxilla 1 with 1 apical
and 1 enlarged medial seta; spines on outer plate
mostly bifid. Inner plate of maxilliped with 1
stout geniculate apical spine and second much
smaller straight spine. Coxae 1-2 poorly setose.
Gill 2 dominant, gill 5 larger than posterior lobe
of coxa 5. Gnathopod 2 lacking surficial button
combs.
Pereopods 3-4 lacking extensive secondary fa-
cial rows of spines on article 5.
Uropods 1-2 lacking huge dorsal setae on pe-
duncles, rami lacking basoventral setae, medial
margins of peduncles not heavily setose.
TYPE-SPECIES.?Dirimus tarlitus, new species.
COMPOSITION.?Unique.
RELATIONSHIP.?This genus forms a close com-
panion to Tottungus by sharing the cowllike head,
baler lobes on maxilliped, the setose dactyls of
pereopods 5-7 (in males as well as females), and
numerous other similarities; but differs in several
important characters. Dirimus has lost the funda-
mental urohaustoriid character of epimeron 2 in
its loss of dominant facial setae and size domi-
nance; it has a reduced row of organized arma-
ments on article 1 of antenna 1 and no combs on
gnathopod 2. Unlike Tottungus, the setae on the
palp of maxilla 1 remain in 1 set, gnathopod 2
retains its typical parachelate condition and coxa
1 is not larger than coxa 2. Other differences
include the further reduction in mandibular rak-
ers, the retention of the normal 2 apical setae on
article 2 of the outer ramus on uropod 3, the
partially articulate inner rami of uropods 1-2, the
thin left lacinia mobilis, the different apical ar-
mament on the inner plate of the maxilliped, and
the much better-developed eyes.
Dirimus tarlitus, new species
FIGURES 52-54
DIAGNOSIS.?With the characters of the genus;
unique feature being loss of setal and size domi-
nance of epimeron 2 and unusually broad and
short telson.
DESCRIPTION OF FEMALE.?Holotype female
"o," 2.22 mm: Head about 1.2 times as long as
wide, rostrum about 10% as long as remainder of
head. Primary flagellum of antenna 1 with 7
articles, formula of medium-length aesthetascs =
0-0-0-2-1-1-0; accessory flagellum 5-articulate.
Facial formula on antenna 1 = 1 setule, 1 seta, 1
setule; on article 4 of antenna 2 in 2 sections of 6
and 9, spines dominant, on article 5 = 1 1 ; article
3 with 1 main seta; flagellum with 6 articles.
Right and left incisors almost simple, formula on
right and left palp article 3 = 3-1-2-1. Inner
plate of maxilla 1 with 1 short apical seta, 1 large
apicomedial seta; outer plate with 9 spines, oc-
casional spine with much more than 2 cusps.
Medial row on inner plate of maxilla 2 with 2
setae. Inner plate of maxilliped with 3 medial
setae; outer plate with 1 apicolateral seta; palp
article 2 with 2 setae in facial row, article 3 with
1 facial seta; basal articles of maxilliped broad-
ened and with weakly developed baler lobe bear-
ing 1 seta (as shown for Tottungus tungus).
Coxa 1 trapezoidal, not larger than coxa 2,
both poorly setulose; coxa 3 with long seta at
posteroventral corner, ventral margin lined with
2 medium setae and short setules, posterior mar-
gin naked.
Palm of gnathopod 2 slightly excavate, with 3
defining teeth (pair medially, 1 tooth laterally),
no minute serrations except for weak row inside
lateral tooth.
Setal and spine formulas on pereopod 3 = 3, 3,
2-1, 3 + 3 + 1 ; pereopod 4 = 5, 3, 2-1, 4 + 3 +
1; posterior margin of article 5 with row of even
serrations. Article 2 of pereopods 5-7 with pro-
gressively weaker setal sizes posteriorly; article 2
of pereopod 5 without mediofacial brush of setae,
dactyl with weak anterior hump-serrations and 2
subapical setae; dactyl of pereopod 6 with 4
subapical setae, of pereopod 7 with 3 subapical
setae.
Peduncular spine formulas of pleopods 1-3 =
2 and 0, 2 and 0, 2 and 0; segmental formulas =
134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Mt \ (IS
FIGURE 52.?Dirimus tarlitus, new species, holotype, female "o," 2.22 mm (Z? gill).
NUMBER 360 135
P5 D7
FIGURE 53.?Dirimus tarlitus, new species, holotype, female "o," 2.22 mm.
KP5
10-5, 9-7, 9-7; basal setal formulas = 6-2-2-2,
2-1-1-1,5-1-1-1.
Epimera 2-3 of similar shape and size, lacking
teeth and facial setae.
Lateral margin on peduncle of uropod 1 with
2 large setae, small setule at corner, 1 apicomedial
seta; of uropod 2 only with apicolateral setule
and 1 long medial seta (probably attached to root
of inner ramus), no dorsal setae; setae of outer
rami = 6 and 4; inner rami short and partially
articulate, bearing 3 and 2 (3?) setae. Apicolateral
corner of peduncle on uropod 3 with 2 setules,
medial margin with 2 small setae, ventromedial
apical row with 4-5 mixed setae, formula on
lateral margin of article 1 on outer ramus =
e-Ee-Ee.
Telson almost twice as wide as long, setal for-
mula - 2P, P/2P, P.
Gland clusters indiscernible.
Female "t," 2.07 mm: Telsonic setal formula
= 2P, 1, latter seta closely contiguous to its op-
posite partner, thus forming central pair (illus.).
DESCRIPTION OF MALE (male "p," 2.01 mm).?
Like female but eyes slightly larger; telsonic setal
formula = 2P, P, and 1 middle seta (illus.).
HOLOTYPE.?QUS, female "o," 2.22 mm (il-
lus.).
TYPE-LOCALITY.?QMW.6383, Middle Banks,
Moreton Bay, Queensland, Sep 1976, S. New-
lands and S. Cook coll., depth and substratum
unknown.
VOUCHER MATERIAL.?Type-locality: female
136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 54.
pW
-Dinmus tarlitus, new species, holotype, female "o," 2.22 mm (p = male "p," 2.01
mm; t = male "t," 2.07 mm).
"m," 2.25 mm; female "n," 2.17 mm; male "p,"
2.01 mm (illus.); male "s," 2.11 mm; QMW.6390:
male "t," 2.07 mm (illus.).
MATERIAL.?QUS, 2 samples (6).
DISTRIBUTION.?Queensland, Moreton Bay,
Middle Banks, 2-12 m, sand.
HAUSTORHDAE
DIAGNOSIS.?Rostrum moderate to weak, head
short, cheek poorly developed. Antenna 1 of haus-
torius form, article 1 short, articles 2-3 progres-
sively shorter, weakly geniculate or not; primary
flagellum elongate, accessory flagellum short. An-
tenna 2 of haustorius form, article 4 expanded,
article 5 shorter and narrower than article 4, these
articles furnished with 1 or more rows of longi-
tudinal facial ornaments, ventral margin of arti-
cle 4 with at least 3 kinds of setae: (1) elongate
plumes; (2) shorter and stiffer glassy spines set in
tandem; (3) bulbar based penicillate setules; fla-
gellum scarcely longer than article 4 of peduncle.
Prebuccal complex massive, upper lip dominant,
epistome scarcely distinct. Mandibles bearing
stubby, poorly toothed incisors; laciniae mobiles
present on both sides and weakly unlike each
NUMBER 360 137
other; rakers poorly serrate, numerous to few;
molar large and very weakly triturative, usually
fuzzy, often furnished with accessory chopper in
form of articulate multifid spine; palp 3-articu-
late, article 3 with numerous outer setae, no setae
of palp article 3 awned, apical group with mostly
simple setae but proximal members bilaterally
serrate, distinctive and disjunct set of inner basal,
short and bilaterally serrate spines also present.
Lower lip with discrete inner lobes, mandibular
extensions of outer lobes absent. Maxilla 1 with
biarticulate palp, inner plate usually triangular
but always evenly and densely setose medially,
lacking any lateral setae; baler lobes often pres-
ent. Maxilla 2 usually with outer plate signifi-
cantly enlarged, inner plate with fully developed
medial row of setae. Maxilliped with expanded
bases bearing baler lobes (usually), otherwise
maxilliped with normally enlarged plates, outer
spinose; palp 3-articulate, article 3 usually genic-
ulate, article 2 expanded.
Coxae 1-4 evenly integrated by stepped en-
largement from coxa 1 onwards, coxa 4 dominant,
coxa 1 not extremely small, coxae 3-4 more or
less weakly produced posteroventrally. Coxal gills
on segments 2-6, brood plates slightly to moder-
ately broadened.
Gnathopods feeble, grossly alike in proportions,
wrists elongate, but gnathopod 1 simple (dactyl
with nail = unusual); gnathopod 2 minutely par-
achelate; article 3 short. Article 5 of pereopods
3-4 broad and deeply lobate, with thick posterior
or facial spines; dactyls of pereopods 3-7 vestigial,
represented by tiny bulb bearing plusetule; per-
eopod 5 of haustorius form, articles 2, 4, and 5
expanded, articles 4-5 with extensive facial rows
of spines; pereopods 6-7 not alike but not of
phoxocephalid form; article 4 on pereopod 6
immensely dominant, article 2 subrectangular;
article 2 on pereopod 7 immensely dominant,
subcircular, articles 4-5 expanded, but together
not as large as article 4 of pereopod 6; pereopods
6-7 occasionally with weakly developed under-
slung articulation.
Peduncles of pleopods not longer than wide,
inner rami not or scarcely inferior, pleopod 2 not
inferior; coupling hooks paired on each pleopod,
inner rami lacking clothespin hooks.
Epimeron 1 well developed; epimeron 3 dom-
inant in size, cuspidation or setation. Urosome
flexing ventrad, urosomite 1 often becoming elon-
gate, extending ventrally, and apicoventrally
toothed. Rami of uropod 1 clavate or rodlike or
weakly styliform, spinose; rami of uropod 2 lin-
guiform or grossly rodlike, setose; uropod 3 with
rodlike rami, outer ramus weakly dominant, 2-
articulate, article 2 elongate, inner and outer rami
and article 1 of outer ramus apically multiarmed.
Telson very short, broad, cleft gaping, base
often constricted.
TYPE GENUS.?Haustorius Miiller.
COMPOSITION.?Lepidactylus Say, Eohaustorius J.
L. Barnard, Protohaustorius Bousfield, Parahaustorius
Bousfield, Neohaustorius Bousfield, Pseudohaustorius
Bousfield, Acanthohaustorius Bousfield. (See Bous-
field, 1965, 1973).
RELATIONSHIP.?Haustoriids can be developed
evolutionarily from a position antecedent both to
the Zobrachoidae-Urohaustoriidae cluster and to
the Phoxocephalopsidae. Haustoriids retain the
primitive epimeral condition of most amphipods
in that epimeron 3 is dominant, whereas the other
groups mentioned have epimeron 2 dominant.
Haustoriids also maintain, like Phoxocephalop-
sidae, the plesiomorphic uropod 1 bearing rod-
styliform rami with spines, quite unlike the lin-
guiform-setose rami of Zobrachoidae-Urohausto-
riidae, which are very unusual and presumably
apomorphic. The rami of uropod 2, however, are
linguiform but the setae are far more numerous
and not as regularly positioned as in the antibo-
real groups. Uropod 3 is distinctive and somewhat
reminiscent of certain Eurasian freshwater gam-
marids.
The presence of baler lobes is a specialization
superimposed on what otherwise are primitive
maxillae in terms of medial setation; but the
maxillipeds have progressed greatly through loss
of the dactyl and geniculate substitution by arti-
cle 3 of dactylar functions. Haustoriids retain
many rakers in primitive forms, though several
apomorphic genera show decline in numbers, as
138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
in Urohaustoriidae. The presence of a special
inner proximal row of spines on palp article 3
may be a carryover from the primitive gammarid
condition; these do not occur in other hausto-
rioids as far as known. ,
Gnathopods are very similar to those of Uro-
haustoriidae but lack surficial button combs on
the faces of article 5 (observation new). The
dactyl of gnathopod 1 bears a nail, a unique
feature. All pereopodal dactyls are truly vestigial
(e.g., as small as can be imagined and be observ-
able). Article 5 of pereopods 3-4 is grossly lobate.
Pereopods 6-7 are of quite distinctive morphol-
ogy in comparison with zobrachoid-urohausto-
riids. The pleopods differ in several ways from
those in antiboreal taxa.
CONDUKHDAE, new family
DIAGNOSIS.?Rostrum well developed, cylindri-
cal, with anteroventral solid keel; head otherwise
elongate, cheek poorly developed. Appendages
fossorial, e.g., furnished with digging spines and
roiling-filtrative setae. Antenna 1 partially of
Urothoe form, article 1 short, articles 2-3 unusually
elongate, geniculate; flagella short, similar and
about as long as article 3 of peduncle. Antenna 2
slender, article 4 elongate, article 5 shorter than
article 4, these articles furnished with long and
medium setae, no rows of facial armaments; fla-
gellum short, subequal to article 5 of peduncle.
Prebuccal complex massive, epistome and upper
lip distinct, latter dominant. Mandibles bearing
incisors, laciniae mobiles, rakers and large weakly
triturative molar; palp 3-articulate. Lower lip
with appressed inner lobes partially fused to-
gether, mandibular lobes of outer lobes well de-
veloped. Maxilla 1 with small, moderately setose
inner plate; outer plate with 6 spines; palp 2-
articulate. Maxilla 2 ordinary, inner plate with
oblique facial row of setae. Maxillipeds with
widened and extended basal parts lacking baler
lobes, outer plate bearing only setae, otherwise
with normally enlarged plates and 4-articulate
palp, dactyl of latter stubby or clavate and mul-
tisetose, lacking nail.
Coxae 1-4 progressively and evenly larger;
coxa 1 small but not strongly hidden by following
coxae; coxa 4 broad, platelike, with weak poster-
odorsal excavation; coxa 5 much shorter than
coxa 4. Goxal gills on segments 2-6, brood plates
unknown.
Gnathopods feeble, grossly alike in proportions
but gnathopod 1 scarcely subchelate; gnathopod
2 minutely parachelate, wrists elongate, article 3
very short. Article 5 of pereopods 3-4 with large
posterior lobe and thick spines; dactyls of pereo-
pods 3-6 vestigial, dactyl of pereopod 7 small;
pereopods 5-7 of similar dominance, article 2
grossly similar in expansion, articles 4-5 broadly
expanded and platelike, bearing facial spine
groups, though weak or absent in several places,
article 6 much narrower and shorter and hanging
as appendage to apex of article 5; article 5 of
pereopod 7 underslung.
Peduncles of pleopods not longer than wide,
rami of pleopods 1 and 3 extremely slender, with
very few articles, those of uropod 2 weakly ex-
panded basally, otherwise similar, with article 1
extremely elongate, only pleopod 2 with coupling
hooks; essentially pleopod 2 dominant.
All epimera distinctly developed, epimeron 3
dominant. Urosome flexed ventralwards (on
death), urosomite 1 greatly elongate by virtue of
huge subpeduncular extension bearing pair of
ventral teeth, urosomites 2-3 equally short; uro-
pods 1-2 each biramous, rami sublanceolate (not
linguiform), spinose (not setose, or setae, if pres-
ent, very short); peduncle of uropod 3 scarcely
elongate, inner ramus vestigial, outer ramus
grossly dominant, article 2 well developed, api-
cally truncate, and multiarmed.
Telson ordinary, slightly longer than wide,
deeply cleft. Sexual dimorphism unknown.
TYPE GENUS.?Condukius, new genus.
COMPOSITION.?Unique.
RELATIONSHIP.?Condukius carries a mixture of
characteristics found in the Haustoriidae, Uro-
thoidae, Pontoporeiidae, and Platyischnopidae.
For example, the rostrum resembles a primordial
concept of the platyischnopid rostrum and the
following appendages are so similar to those of
NUMBER 360 139
platyischnopids that a widened diagnosis could
admit Condukius: pereopods 5-7, uropods 1-3,
telson, epimera, lower lip, and maxilliped palp
article 4. However, the following characteristics
of Condukius are foreign to platyischnopids: the
simple gnathopod 1, the general shape of gnath-
opod 2, and in most respects the parts of the
mandibles, maxilla 1, maxilliped and pereopods
3-4.
The strange apical setae on the wrists of the
gnathopods are also characteristic of several pla-
tyischnopids and alien to the other groups.
Condukius differs from both Urothoidae and
Haustoriidae in the shape and spination pattern
of pereopod 5 and in the rostrum, but pereopods
3-4 have features of the Haustoriidae. The outer
plate of the maxilliped, in its lack of thick spines,
resembles that of the Haustoriidae. Condukius dif-
fers from the Haustoriidae in the presence of
mandibular lobes on the lower lip, the presence
of maxillipedal palp article 4, an elongate telson,
relatively normal spination on uropod 2 and dis-
tinctive antennae.
Condukius might be included within the Ponto-
poreiidae because of resemblances in telson, lower
lip, uropods 1-2, and gnathopods, but differs in
the rostral shape and antenna 1.
Provisionally, Condukius may be assigned to
Platyischnopidae on the basis of rostrum and
characteristic setae of the gnathopods but segre-
gation in a distinct family on the basis of shapes
in articles 4-5 of pereopods 6-7 (like Haustori-
idae); shape of pereopods 3-4 (like Haustoriidae);
the absence of thick spines on the outer plate of
the maxilliped (like Haustoriidae); and the unu-
sual mixture of pleopods; is a good alternative.
Condukius shares some of the unusual appear-
ances found in Cunicus, now assigned to the Uro-
thoidae. These similarities occur especially in an-
tennae and mouthparts and the slight propensity
in Cunicus to the development of a protruding
rostral area; but Cunicus differs from Condukius
greatly in that (1) the head is readily identifiable
with the Urothoe-likc head, with large cheek, and
the antennae are attached at the apex of the false
rostrum; (2) pereopod 5 is of the haustorius form;
pereopods 6-7 are of the urohaustorius form; (3)
the rami of uropods 1-2 are absent, but the
remaining peduncle is lamelliform and strongly
setose.
Condukius, new genus
DIAGNOSIS.?With the characters of the family,
especially notable being: Platyischnopid-like ros-
trum, thin geniculate antennae lacking organized
facial armaments, with elongate setae, articles
2-3 of antenna 1 elongate; relatively ordinary
mandibles, coxae, and telson; feeble gnathopods
with elongate wrists, if chelate not strongly so;
vestigial dactyls on pereopods 3-7; gross lobation
on article 5 of pereopods 3-4; evenness of pereo-
pods 5-7, all with expanded articles 2, 4, and 5
bearing facial spines, pereopod 5 not of the haus-
torius form; unusual pleopods; ordinary uropods
1-2; niphargid-like uropod 3; flexed urosome
with greatly attenuate urosomite 1.
TYPE-SPECIES.?Condukius karkan, new species.
COMPOSITION.?Unique.
Condukius karkan, new species
Figures 55-58
DIAGNOSIS.?With the characters of the genus
and family.
DESCRIPTION.?Body long and narrow, as in
Platyischnopidae. Head 17% as long as total an-
imal, narrow and elongate, not tall, with platyis-
chnopid-like rostrum extending anteriorly; eyes
indiscernible; antenna 1 attached in sinus behind
rostrum, ventroanterior margin of head lacking
sinus for insertion of antenna 2, cheek not ven-
trally extended.
Antenna 1 shorter than antenna 2, article 1 as
long as wide, dorsofacial surface with 2 large
setae; article 2 nearly 1.7 times as long as article
1, slender, with 2 large setae; article 3 slightly
shorter than article 2 but elongate (in context to
many similar genera), with midventral but facial
circlet of 5 setae and 1 apicodorsal seta; flagella
almost equally extended but only 1.25 times as
140 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 55.?Condukius karkan, new species, holotype, male "a," 8.41 mm (Y = pleopod).
NUMBER 360 141
Mt
HR3
FIGURE 56.?Condukius karkan, new species, holotype, male "a," 8.41 mm.
long as article 3 of peduncle, primary flagellum
6-articulate, penultimate article short, other elon-
gate, bearing pairs of hoodscales apically except
for ultimate article forming elongate tube; acces-
sory flagellum 7-articulate, otherwise similar but
penultimate article not shortened. Article 3 of
antenna 2 with 2 setae; article 4 slender, ventral
margin closely lined with elongate setae, dorsal
margin with fewer and shorter setae, face with 2
setae at about M. 60; article 5 much shorter than
article 4, ventral setae irregular and shortened,
dorsal setae of medium size, apical face with
several setae; flagellum almost as long as article
5 of peduncle, similar to flagella of antenna 1
(also bearing hoods and ultimate article tubelike).
Mandibular incisors short, right weakly bifid,
left weakly trifid; right and left laciniae mobiles
not grossly dissimilar, of moderate thickness, right
bifid, with 3 other humps superimposed, left
weakly bifid; rakers expanded and serrate, right
with 5 plus 2 rudimentaries, plus intercalated
plusetules, left with 6 plus 2 rudimentaries and
plusetules; molars large, elongate, weakly tritur-
ative, bearing accessory choppers with 5 cusps;
palps ordinary, all setae of article 2 concentrated
on midmedial margin; article 3 formulas not of
Urohaustoriidae format, apex of palp with defi-
nite bevel and all setae concentrated on this inner
bevel, setae simple but apically hooked. Lower
lip lacking cones.
142 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
>ZG2
FIGURE 57.?Condukius karkan, new species, holotype, male "a," 8.41 mm (Z ? gill).
NUMBER 360 143
W
FIGURE 58.?Condukius karkan, new species, holotype, male "a," 8.41 mm (Y = pleopod).
144 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Inner plate of maxilla 1 with 2 apicolateral
setules, 1 large apical seta and 2 similar medial
setae all in tandem: outer plate tapering apically,
bearing only 6 spines, 2 of these bifid; palp biar-
ticulate, article 1 short, article 2 elongate and
broadly expanded (when flattened), multisetose
apically, medially, and facially. Inner plate of
maxilla 2 narrower and slightly shorter than outer
plate, with mediofacial oblique row of 4 setae.
Inner plate of maxilliped with 3 stout apical
spines, oblique facial row of 5 setae; outer plate
with medial and apical setae, no spines; palp
attached geniculately to outer plate article, article
2 moderately expanded and densely setose medi-
ally and facially, article 3 attached to article 2 by
narrowed base, then medially expanded and se-
tose medially and facially, dactyl very small,
clavate, with 3-4 apical setae.
Gnathopods densely setose, dactyls small, weak
setal brush present on article 3. Article 4 of
pereopod 3 with 4 large anterior setae, of pereo-
pod 4 with only 1 seta, posterior margins respec-
tively with 1-1 and 2-3; lobe of article 5 unor-
namented, with 2 apposing rows of spines, about
4 in each row (with aberration of seta replacing
spine); article 6 elongate, beveled ventroapex
with 2 rows of spines, 5-4 and 6-4 (perhaps
actually 5-4-1 but single midspine difficult to
interpret); dactyls vestigial, about as large as
basal attachment circle of adjacent spine, bearing
large setule. Coxa 5 ordinarily bilobed; coxa 6
with small anterior and deep posterior lobe; coxa
7 extremely short but wide; article 2 of pereopods
5-7 almost naked posteriorly; otherwise see illus-
trations.
Pleopods unusual (Figure 58: Fl, 2, 3); pleo-
pods 1 and 3 lacking coupling hooks, only pleo-
pod 2 attached to sternum in normal situation
with peduncles set transversely, other pleopods
with peduncles tilted inward, thus when flattened
and illustrated rami appearing superimposed, as
attached to sterna with rami curling backwards;
ramal bases and peduncle of pleopod 2 with
dense, filamentous setulation, other pleopods
lacking this armament; segmental formulas =
4_3; 4_3 and 4-3 (inner with weakly indicated
fourth article at apex of article 1).
Both epimera 1-2 with sharp, hooked, poster-
oventral tooth, posterior margin convex, armed
with 2 and 4 setules; ventral margin of epimeron
1 with 5 setae; posterior part of ventral margin
on epimeron 2 with 2 spines; epimeron 2 much
larger than epimeron 1, each with weak to strong
facial ridge. Epimeron 3 overall quadrate and
somewhat brokenly but normally separated dor-
sally from segment; posterior and posterodorsal
margin with 6 setae, ventralmost largest, ventral
margin naked, face with horizontal row of 3 setae
attached to scallop-ridge.
Urosomite 1 grossly extended ventrally, with 2
clothespinlike ventral teeth side by side (and
asymmetrical); peduncle of uropod 1 with 3-4
ventrolateral setae, dorsally with 2-3 setae near
apex and apical spine, medial apex with spine;
outer ramus with 1 small dorsal seta near base,
apex with 5 widely spread long to medium spines,
inner ramus with 4 similar spines and 2 in tandem
more proximally; peduncle of uropod 2 with 4
dorsolateral spines, apicalmost stoutest, medial
margin with 4 spine-setae, 3 of these crowded
distally, outer ramus with 2 middorsal spines and
group of 5 long to medium apical and apicome-
dial spines, inner ramus with 2 midmedial spine-
setae, apex with 3 shorter spine-setae.
Uropod 3 attached to urosomite 3 and outer
ramus to peduncle apparently with ontogenetic
dorsal face pointing laterally; inner ramus
scarcely detectable; peduncle and article 1 of
outer ramus with faciolateral groups of spines,
new dorsal margins spinose and article 1 also with
large setae, article 2 smaller than article 1, apex
truncate and multispinose, bearing also 1 plu-
mose seta. Telson with tapered apices, at begin-
ning of taper each armed with transverse lateral
set of 3 setae, lateralmost large and elongate,
other medial members much smaller; basolateral
margins naked.
ILLUSTRATIONS.?Armaments added in dots on
uropods 1-2 representing those added from op-
posite member; pereopod 5 illustrated as compos-
ite of left articles 1-4 and converted right 5-7,
NUMBER 360 145
pereopod 6 with right and left armaments com- HOLOTYPE.?NMV, male "a," 8.41 mm.
bined together for maximum count; gills 2-4 Unique.
drawn at same magnification as those attached TYPE-LOCALITY.?WPBES 1745/1 Western
to pereopods 5-6; pleopods drawn from left pos- p o r t Victoria, 25 Nov 1974, 13 m, coarse-me-
tenor view, dotted setae showing additional mem- i- ,
bers from right side.
Literature Cited
Barnard, J.L.
1961. Gammaridean Amphipoda from Depths of 400 to
6000 Meters. In Galathea Report, 5:23-128, 83 fig-
ures.
1972. Gammaridean Amphipoda of Australia, Part I.
Smithsonian Contributions to Zoology, 103: 333 pages,
194 figures.
Barnard, J.L., and C M . Barnard
1982. The Genus Rhepoxynius (Crustacea: Amphipoda:
Phoxocephalidae). Smithsonian Contributions to Zool-
ogy, 357: 49 pages, 6 figures.
In press. The Freshwater Gammaridea (Amphipoda) of
the World. Smithsonian Contributions to Zoology.
Barnard, J.L., and M.M. Drummond
1978. Gammaridean Amphipoda of Australia, Part III:
The Phoxocephalidae. Smithsonian Contributions to
Zoology, 245: 551 pages, 269 figures.
1979. Gammaridean Amphipoda of Australia, Part IV.
Smithsonian Contributions to Zoology, 269: 69 pages,
38 figures.
Bousfield, E.L.
1965. Haustoriidae of New England (Crustacea: Am-
phipoda). Proceedings of the United States National
Museum, 117:159-240,31 figures.
1973. Shallow-Water Gammaridean Amphipoda of New Eng-
land. Ithaca: Cornell University Press.
Dana, J.D.
1853. Crustacea, Part II. In United States Exploring Expe-
dition, 14:689-1618, 96 plates.
Fearn-Wannan, H.J.
1968. Littoral Amphipoda of Victoria, Part I. Proceedings
of the Royal Society of Victoria, new series, 81:31-58,
18 figures.
Schellenberg, A.
1931. Gammariden und Caprelliden des Magellange-
bietes, Siidgeorgiens und der Westantarktis. In
Further Zoological Results of the Swedish Antarctic Ex-
pedition, 1901-1903, 2(6): 1-290, 136 figures,
plate 1.
Sheard, K.
1936. Amphipoda from a Southern Australian Reef,
Part 1. Records of the South Australian Museum,
5:445-455, figures 1-4.
Sivaprakasam, T.E.
1970. Amphipoda from the East Coast of India, 2:
Gammaridea and Caprellidea. Journal of the
Bombay Natural History Society, 67:153-170,
figures 13-18.
Stebbing, T.R.R.
1906. Amphipoda, I: Gammaridae. Das Tierreich, 21: 806
pages, 127 figures.
Stephensen, K.
1947. Tanaidacea, Isopoda, Amphipoda, and
Pyconogonida. In Scientific Results of the
Norwegian Antarctic Expeditions, 1927-1928. Det
Norske Videnskaps-Akademi i Oslo, 27:1-90,
figures 1-26.
Stephenson, W., S.D. Cook, and SJ . Nedlands
1978. Macrobenthos of the Middle Banks Area of
Moreton Bay. Memoirs of the Queensland Museum,
18:185-212.
146
Index
(Initial page numbers of principal accounts in italics)
Acanthohaustorius, 5, 137
Amphiporeia, 5, 6
Argissa, 5
Argissidae, 5
barinius, Tuldarus, 119, 123
Bathyporeia, 5, 6, 7, 9, 14
buchalius, Bumeralius, 28
Bumeralius, 6, 9, 18, 19, 27, 37, 48, 107
buchalius, 28
cangellus, Tuldarus, 119, 126
canguro, Zobracho, 20
Carangolia, 6, 7, 15, 37
Cardenio, 5
Compactogammarinae, 11
Condukiidae, 11, 12, 14, 138
Condukius, 138, 139
karkan, 139
Cunicus, 6, 7, 10, 15, 37, 139
deceptionis, Phoxocephalopsis, 16
Dexaminidae, 5
Dirimus, 6, 9, 46, 47, 132
tarlitus, 133
elegans, Urothoe, 7
Eohaustorius, 5, 137
garbaius, Gheegerus, 107, 108
Gheegerus, 6, 9, 46, 47, 107
garbaius, 107, 108
gunni, Urohaustorius, 48, 50, 52, 96, 103, 107
halei, Urohaustorius, 48, 50, 52, 89, 97, 100, 102, 106, 107
Harpinia, 5
Haustoriidae, 5, 9, 11, 12, 136
Haustoriinae, 5
Haustoriopsis, 5
Haustorius, 5, 6, 137
Indischnopus, 14
Japara, 4
karkan, Condukius, 139
Kotla, 7
Lepidactylus, 5
merkanius, Urohaustorius, 48, 49, 50, 51, 53, 59, 60, 64, 70,
73, 74, 76, 80, 83, 84
metungi, Urohaustorius, 48, 49, 50, 52, 85, 89, 94, 95, 96, 97,
103, 106, 107
Narunius, 6, 9, 46, 47, 107, / / / , 119
tallerkus, 97, 112
Neohaustorius, 5, 137
Oedicerotidae, 11
Parahaustorius, 5
parnggius, Urohaustorius, 48, 49, 50, 51, 52, 53, 64, 68, 69,
70, 73, 74, 80, 83, 84
pentinus, Urohaustorius, 48, 49, 50, 51, 52, 60, 61, 69, 72,
73, 76, 84, 85, 97
perkeus, Urohaustorius, 48, 50, 51, 52, 80, 85, 112
Phoxocephalidae, 5, 9, 11, 72, 14
Phoxocephalinae, 14
Phoxocephalopsidae, 11, 12, 15
Phoxocephalopsis, 5, 6, 10, 16
deceptionis, 16
zimmeri, 16
Phoxocephalus, 5
Platyischnopidae, 5, 11, 12, 13
Platyischnopus, 5, 14
Pontogammaridae, 7, 11
Pontoporeia, 5, 6, 7
Pontoporeiidae, 7, 11
Pontoporeiinae, 5
Prantinus, 6, 9, 10, 11, 15, 18, 36, 37, 46, 48
talanggi, 37, 38
Priscillina, 5, 6, 7
Protohaustorius, 5, 137
Pseudohaustorius, 5, 137
pulcus, morph 872, Urohaustorius, 49, 59
pulcus, Urohaustorius, 48, 49, 50, 51, 52, 53, 59, 60, 61, 63,
68, 69, 70, 72, 73, 76, 80, 84, 85, 89, 111, 112, 117
species M, Urohaustorius, 49, 51, 53, 84
Synchelidium, 11
Synopiidae, 5
talanggi, Prantinus, 37, 38
tallerkus, Narunius, 97, 112
tarlitus, Dirimus, 133
Tironidae, 5
Tittakunara, 14
Tomituka, 14
Tottungus, 6, 9, 10, 46, 47, 107, 126, 133
tungus, 127, 133
Tuldarus, 6, 9, 46, 47. 107, 118, 127, 132
barinius, 119, 123
cangellus. 119. 126
tungus, Tottungus, 127, 133
Urohaustoriidae, 9, 11, 12, 16, 45, 137
Urohaustorius, 5, 6, 9, 19. 20. 25, 27, 28. 37, 46. 47, 69, 84,
107, 119, 131
gunni, 48, 50, 52, 96. 103, 107
halei, 48, 50, 52, 89, 97, 100, 102, 106, 107
147
148 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
merkanius 48, 49, 50, 51, 53, 59, 60, 64, 70, 73, 74, 76, 80,
83,84
metungi, 48, 49. 50. 52, 85, 89, 94, 95, 96, 97, 103, 106,
107
parnggius, 48. 49. 50, 51, 52, 53, 64, 68, 69, 70, 73, 74, 80,
83, 84
pentinus, 48, 49, 50, 51, 52. 60, 61, 69, 72, 73, 76, 84, 85,
97
perkeus, 48, 50, 51, 52, 80, 85, 112
pulcus, 48, 49, 50, 51, 52, 53, 59, 60, 61, 63, 68, 69, 70, 72,
73, 76, 80, 84, 85,89, 111, 112, 117
pulcus, morph 872, 49, 59
species M, 49, 51, 53, 84
urungari, 48, 49, 50, 51, 53, 70, 77, 79, 80
vercoi, 48, 97
wingaro, 48. 49, 50, 51, 52, 60, 70, 76, 80, 83, 84
yurrus, 48, 49, 50, 51, 52, 53, 70, 76, 80, 81, 84, 85
Urothoe, 5, 6, 7, 9, 15, 37, 138, 139
elegans, 7
Urothoidae, 11, 12, 14
Urothoides, 5, 6, 7, 15, 37
urungari, Urohaustorius, 48. 49, 50, 51, 52, 53, 70, 77, 79, 80
vercoi, Urohaustorius, 48, 97
wingaro, Urohaustorius, 48, 49, 50, 51, 52, 60, 70, 76, 80, 83,
84
Yammacoona, 7
Yurrokus, 14
yurrus, Urohaustorius, 48, 49, 50, 51, 52, 53, 70, 76, 80, 81,
84, 85
zimmeri, Phoxocephalopsis, 16
Zobracho, 6, 9, 10, 11, 15, 16, 18, 19, 28, 37, 48
canguro, 20
Zobrachoidae, 11, 12, 16, 18, 137
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for volume/number/page citations: "10(2):5-9." For alinement and arrangement of
elements, follow the format of the series for which the manuscript is intended.
Legends for illustrations must not be attached to the art nor included within the text but
must be submitted at the end of the manuscript?with as many legends typed, double-
spaced, to a page as convenient.
Illustrations must not be included within the manuscript but must be submitted sepa-
rately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.)
can be intermixed throughout the printed text. They should be termed Figures and should
be numbered consecutively. If several "figures" are treated as components of a single larger
figure, they should be designated by lowercase italic letters (underlined in copy) on the illus-
tration, in the legend, and in text references: "Figure 9JD." If illustrations are intended to be
printed separately on coated stock following the text, they should be termed Plates and any
components should be lettered as in figures: "Plate 9b_." Keys to any symbols within an
illustration should appear on the art and not in the legend.
A few points of style: (1) Do not use periods after such abbreviations as "mm, ft,
yds, USNM, NNE, AM, BC." (2) Use hyphens in spelled-out fractions: "two-thirds." (3)
Spell out numbers "one" through "nine" in expository text, but use numerals in all other
cases if possible. (4) Use the metric system of measurement, where possible, instead of
the English system. (5) Use the decimal system, where possible, in place of fractions.
(6) Use day/month/year sequence for dates: "9 April 1976." (7) For months in tabular list-
ings or data sections, use three-letter abbreviations with no periods: "Jan, Mar, Jun," etc.
Arrange and paginate sequentially EVERY sheet of manuscript?including ALL front
matter and ALL legends, etc., at the back of the text?in the following order: (1) title page,
(2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes,
(7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends.
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