PROC. BIOL. Soc. WASH.
102(4), 1989, pp. 960-967
NEW GENERA IN THE THALASSINIDEAN
FAMILIES CALOCARIDIDAE AND AXIIDAE
(CRUSTACEA: DECAPODA)
Abstract. - The family Calocarididae Ortmann, 1891, is resurrected and re-
diagnosed. Three characters distinguish members of the family: hermaphro-
ditism, eye reduction, and second pleopods having enlarged appendices mas-
culinae along with loss of the distal endopod. The first pleopods are spatulate
and somewhat plow-shaped, a character also seen in some axiids. Four genera
are included in the Calocarididae: Calastacus, Calocaris, Callistocaris, and
Lophaxius, the latter two being described as new. Two new genera in the Axiidae
are described: Posthonocaris and Sakaiocaris. A scheme for the derivation of
the Calocarididae from more generalized axiids such as the two newly-described
genera, is proposed.
The families and many of the genera of
the Thalassinidea are extremely poorly de-
fined, with little sense of phylogeny in the
currently accepted classification. The Axi-
idae in particular contains several more or
less well diagnosed and probably polyphy-
letic genera. In the course of a phylogenetic
study of the thalassinideans, it became clear
that a group of four genera ofaxiids were
more closely related to each other than to
the rest of the family. Almost 100years ago,
Ortmann (1891), placed one of these genera
in a separate family, the Calocaridae, but
this taxon received no recognition and has
not been used in the last 50 years. This fam-
ily is now resurrected and rediagnosed for
this group of four probably monophyletic
genera.
Calocarididae Ortmann, 1891
Calocaridae Ortmann, 1891:47, 50.-Ste-
phensen, 1910:75, 77, 189.- Runnstr6m,
1925:14.
Type genus. - Calocaris Bell, 1853.
Diagnosis. -Hermaphroditic, generally
deepwater forms, with eyes showing some
reduction and loss of both pigment and om-
matidial facets. Linea thalassinica absent.
Maxilla 2 scaphognathite bearing spinulose
whip. Maxilliped 3 bearing dentate ischial
crest. Pereopods lacking exopods. Pereo-
pods 1 and 2 chelate. Pleopodal rami elon-
gate-slender. Pleopod 1 always present, of
2 articles, directed anteromesially along
posterior thoracic sternites, with distal ar-
ticle broadened and flattened, and bearing
proximo mesial cluster of hooks (fused ap-
pendix interna). Pleopod 2 present, showing
loss of distal endopod, enlargement of ap-
pendix masculina, and eventual fusion of
appendix interna with appendix masculina.
Uropodal outer ramus with transverse su-
ture.
Genera. - Calastacus Faxon, 1893; Cal-
listocaris, new genus; Calocaris Bell, 1853;
Lophaxius, new genus.
Remarks. -Ortmann (1891) designated
the new family Calocaridae, characterized
by four features: rostrum flat, triangular;
pleopodal rami narrow; outer uropodal ra-
mus with suture; podobranchs and mas-
tigobranchs present on pereopods. Ort-
mann further suggested that eventually two
subfamilies, the Calocarinae and the Lao-
midiinae, would need to be distinguished.
Stephensen (1910), in a rather obscure semi-
popular handbook, used the name Calocar-
idae as a family of the Anomura, though not
designating an author. His diagnosis, loose-
ly translated, reads: "The large chelae have
straight long fingers, but no tubercles. Two
pairs oflegs have chelae. Suture across outer
branches ofuropod. (*Footnote-By a mis-
fortune this suture was dropped from fig.
51). Only one genus and species with us"
[presumably meaning 'in Denmark']. Fol-
lowing this diagnosis, is a short discussion
and figureof Calocaris macandreae Bell.The
only other use of the name Calocaridae I
have been able to trace, is that of Runn-
strom (1925).This family name (in corrected
form), is now resurrected, and its diagnosis
expanded, to include several related genera,
but not the genera of the Laomidiidae, which
are markedly different.
Calastacus Faxon, 1893
Calastacus Faxon, 1893:194.-Borradaile,
1903:539.-de Man, 1925:8, 116.-Balss,
1957:1580.-de Saint Laurent, 1972:353,
354.
Type species. -By monotypy, Calastacus
stilirostris Faxon, 1893:194.
Diagnosis. -Hermaphroditic. Carapace:
supraocular spine present; post-cervical ca-
rina and spines lacking; rostrum at lower
level than anterior carapace; rostral margins
unarmed (except for supraocular spine);
median carina entire; submedian carina
lacking; lateral carina present only as short
posterior extension of lateral rostral mar-
gins. Eye lacking pigment, stalk equal to or
shorter than cornea. Antennal acicle a slen-
der, elongate spike.
Maxillipeds: exopods on 1-3; epipods on
1-3, with small podobranch on 3; 2 arthro-
branchs on 3.
Pereopods: lacking exopods; epipods on
1-4, small podobranchs on 1-3; 2 arthro-
branchs on 1-4; pleurobranchs lacking; pro-
podi and dactyli of 3-5 cylindrical, not ex-
panded; pereopod 1 chelipeds slightly
asymmetrical, not sexually dimorphic; pe-
reopod 2 chelate; pereopods 3-5, dactyli
simple.
Pleopods: rami elongate-slender; appen-
dix intern a present on 2-5; pleopod 1 of 2
articles, distal article broad, plow-shaped,
appendix interna represented by small me-
sial lobe bearing hooks; pIeopod 2 with small
appendix interna fused with basal article of
appendix masculina; both articles of latter
with double row of setae on mesial margin.
Uropod: outer ramus with transverse
dentate suture.
Telson: lacking dorsal spines, longer than
wide.
Species.-
Calastacus laevis de Saint Laurent, 1972.
Bay of Biscay, north-east Atlantic, 950-
1000 m.
Calastacus stilirostris Faxon, 1893. Pacific
Mexico, 1208 m.
Callistocaris, new genus
Type species. -By present designation,
Calocaris alcocki (McArdle, 1900).
Etymology. - The generic name is de-
rived from the Greek "kallistos" - most
beautiful, plus "karis" -a shrimp. Gender:
feminine.
Diagnosis. - Hermaphroditic. Carapace:
strong supraocular spine present; postcer-
vical carina and spines lacking; rostrum set
slightly lower than anterior carapace; rostral
margins armed; median carina entire; sub-
median carina entire; lateral carina entire
posterior to supraocu1ar spine. Eye lacking
pigment, anteriorly flattened, with mesio-
distal tubercle; eye and stalk not differen-
tiated. Antennal acicle fairly well devel-
oped, spike-like, but considerably less than
half length of peduncle article 4.
Maxillipeds: exopods and epipods (no
podobranchs) on 1-3; 2 arthrobranchs on 3.
Pereopods: exopods lacking; epipods on
1-4, podobranchs lacking; 2 arthrobranchs
on 1-4; pleurobranchs lacking; propodi and
dactyli of 3-5 not expanded; pereopod 1
chelae symmetrical, lacking gape between
fingers; pereopod 2 chelate; dactyli of 3-5
simple. Branchiae simple, lacking pinnae.
Pleopods: rami elongate-slender, appen-
dix interna lacking on 3-5, pleopod 1 uni-
ramous, biarticulate, distal article bilobed,
with small mesial patch of hooks; pleopod
2 appendix masculina with indication offu-
sion of 2 articles, with double row of spines
along mesial margin, and with appendix in-
terna fused basally.
Dropod: outer ramus with transverse non-
dentate suture.
Telson: longer than wide, lacking dorsal
spmes.
Species.-
Callistocaris aberrans (Bouvier, 1905). Off
St. Lucia, Lesser Antilles, 809 m.
Callistocaris alcocki (McArdle, 1900). Bay
of Bengal, 992 m.
Callistocaris cf.alcocki (McArdle, 1900).SW
Indian Ocean, 1000 m.
Calocaris Bell, 1853
Calocaris Bell, 1853:231. -Ortmann, 1891:
50, pI. 1, fig. 5d-i.-Borradai1e, 1903:
539.-deMan, 1925:7, 114.-Balss, 1957:
1580.-de Saint Laurent, 1972:353, 354.
Type species. -By monotypy, Calocaris
macandreae Bell, 1853:233.
Diagnosis. -Hermaphroditic. Carapace:
supraocular spine part oflateral rostral spine
series; post-cervical carina entire; rostrum
at slightly lower level than anterior cara-
pace; rostral margins armed; median carina
entire; submedian carina absent; lateral ca-
rina armed. Eye lacking pigment, stalk and
cornea not differentiated; anteriorly flat-
tened and contiguous along midline. An-
tenna1 acicle reduced to barely visible scale.
Maxillipeds: exopods and epipods on 1-
3; reduced podobranch on 2 and 3; 2 ar-
throbranchs on 3.
Pereopods: lacking exopods; epipods on
1-4, small podobranchs on 1-3; 2 arthro-
branchs on 1-4; pleurobranchs lacking; pro-
podi and dactyli of 3-5 not expanded; pere-
opod 1, chelae subsimilar, symmetrical, not
sexually dimorphic, broad gape between fin-
gers; pereopod 2 chelate; pereopods 3-5
dactyli simple.
Pleopods: pleopod 1 of 2 articles, distal
article expanded, lobed, with patch of hooks
on small mesiodistal lobe; pleopod 2, en-
dopod lacking distal portion, appendix
masculina slender, mesially setose, of single
article, appendix intern a articulating at its
base.
Dropod: outer ramus with transverse non-
dentate suture.
Telson: longer than wide, with or lacking
two rows of submedian non-articulating
dorsal spines.
Species. -
Calocaris barnardi Stebbing, 1914. Off Sal-
danha Bay, South Africa, 89-180 m. Off
Namibia, 338 m.
Calocaris granulosus Grebenyuk, 1975.Gulf
of Alaska.
Calocaris macandreae Bell, 1853. Mediter-
ranean; North-east Atlantic, 25-1072 m.
Calocaris sp. Indian 'form' of C. macan-
dreae (see Alcock, 1901): Bay of Bengal,
Arabian Sea.
Calocaris templemani Squires, 1965.
Northwest Atlantic, Newfoundland, Gulf
of Maine, 260 m.
Lophaxius, new genus
Type species. -By present designation,
Lophaxius rathbunae, new species (=Ca-
lastacus investigatoris Rathbun, 1904, non
Anderson, 1896).
Etymology. - The generic name is de-
rived from the Greek 'lophos,' a crest or
mane, referring to the post-cervical mid-
dorsal ridge of the carapace, plus the fre-
quently-used 'axius.' Gender: masculine.
Diagnosis. -Hermaphroditic. Carapace:
supraocular spine (part of lateral rostral se-
ries) present; post-cervical carina with ir-
regular tubercles present; rostrum at slightly
lower level than anterior carapace; rostral
margins armed; median carina entire; sub-
median carina absent; lateral carina only
extending short distance posterior to ros-
trum, with one or two spines. Cornea un-
pigmented, not flattened; stalk subequal to
cornea in length. Antennal acicle short.
Maxillipeds 1-3 with exopods and epipods;
large podobranch on 2 and 3; 2 arthro-
branchs on 3.
Pereopods: exopods lacking; epipods on
1-4, with large podobranch on 1-3; 2 ar-
throbranchs on 1-4; pleurobranchs lacking;
pereopod 1 symmetrical, fingers of chelae
gaping basally; pereopod 2 subchelate; dac-
tyli of 3-5 simple.
Pleopods: rami elongate-slender; appen-
dix interna present on 2-5; pleopod 1 of 2
articles, distal article spatulate, with small
mesial patch of hooks; pieopod 2, appendix
masculina slender, tapering, setose, with ap-
pendix interna articulating at its base.
Uropod: outer ramus with transverse non-
dentate suture.
Telson: longer than wide, with dorsal non-
articulating spines.
Species.-
Lophaxius investigatoris (Anderson, 1896).
Arabian Sea, 1733 m.
Lophaxius rathbunae, new species. North-
eastern Pacific, Alaska to California, 549-
1190 m.
Remarks. - Lophaxius resembles Calas-
tacus in having non-pigmented eyes, and
with the cornea not flattened as in Calo-
caris. It differs from typical Calastacus in
having spines on the rostrum, pleopod 1not
as broadly plow-shaped, the appendix mas-
culina of pleopod 2 not as modified, the
appendix intern a free, in having a well de-
veloped postcervical carina, and in having
the chela of pereopod 1with a gap between
fingers as in Calocaris.
Lophaxius rathbunae, new species
Calastacus investigatoris Rathbun, 1904:
151, non Anderson, 1896.-Schmitt,
1921:112, fig. 75, non Anderson, 1896.
Diagnosis. -Carapace, palm of first che-
la, abdominal somites, uropods, and telson
tuberculate. Merus of pereopod Iwith about
10 spines on anterior (upper) margin, eight
spines on posterior (lower) margin. Outer
uropodal ramus with six spines on outer
margin; inner uropodal ramus with two or
three spines on outer margin.
Material.-Syntypes, USNM 28316, cl
19.5 mm, Albatross sta 3347, off Cascade
Head, Oregon, 631 m.-USNM 28317, cl
19.1mm, Albatross sta 3210, south of Dan-
nakh Islands, Alaska, 884 m. - USNM
28318, ovig. cl 18.4 mm, 17.1 mm, Alba-
tross sta 2928, off San Diego, California,
763 m.-USNM 155734, cl17.1 mm,AI-
batross sta 4352, off San Diego, California,
979-1005 m.
Remarks. - The species Calastacus inves-
tigatoris Anderson, 1896, from 1733 m in
the Arabian Sea is, from the description and
figure (Alcock & Anderson 1896:pl. 25, fig.
1) remarkably similar to the north-eastern
Pacific species, but differs in having weaker
spination on the anterior and posterior mar-
gins of the merus of the first chelipeds, and
in lacking marginal teeth on the inner uro-
podal ramus. No doubt further differences
would be apparent, were material of the In-
dian Ocean species available. Indeed, Rath-
bun (1904: 151) mentioned that not all the
eastern Pacific specimens agreed with Al-
cock's description; she also noted some
variability in the specimens at her disposal.
Given the vast distance between the Indian
and Pacific Ocean records, and the differ-
ences noted, the two species cannot be re-
garded as conspecific.
Key to genera of the Calocarididae
1. Post-cervical carina or ridge present
on carapace 2
- Carapace lacking post-cervical ca-
rina 3
2. Eyes flattened, mesially contiguous
.......................... Calocaris
- Eyes rounded, not mesially contig-
uous Lophaxius
3. Eyes rounded; appendix masculina
mesially setose Calastacus
- Eyes flattened; appendix masculina
mesially spinose Callistocaris
Axiidae Huxley, 1878
Posthonocaris, new genus
Type species. - By present designation,
Axius rudis Rathbun, 1906.
Etymology. - The generic name is a com-
bination of the Greek 'posthon' -one hav-
ing a large penis (referring to the large ap-
pendix masculina), plus 'karis' -a shrimp.
Gender: feminine.
Diagnosis. -Gonochoristic, but with her-
maphroditic forms occurring in the popu-
lation. Carapace: cervical groove present;
postcervical carina and spines absent; ros-
tral margins armed; median carina entire;
submedian carina dentate; lateral carina en-
tire (apart from supraocular spine). Eyeswell
pigmented; eyestalk rounded, longer than or
subequal to cornea. Antennal acicle a well
developed spike.
Maxillipeds: exopods on 1-3; epipod
present on 1 and 2; 3 with epipod plus po-
dobranch, two arthrobranchs.
Pereopods: exopods absent; pereopods 1-
3 with epipod plus podobranch; pereopod
4 with epipod only; two arthrobranchs on
1-4; one pleurobranch on 2-4. Pereopod 1
chelae asymmetrical. Pereopods 3-5, dac-
tyli simple.
Pleopods: Rami elongate-slender. Pleo-
pod 1 in female slender-elongate, of 2 ar-
ticles, distal article bearing marginal setae;
pleopod 1 of male or hermaphrodite spat-
ulate, of 2 articles, distal article bearing
proximomesial clump of hooks; pleopod 2
of male or hermaphrodite with distal setose
portion of endopod somewhat reduced, ap-
pendix masculina and appendix interna ar-
ticulating at about midlength of endopod,
appendix masculina elongate, setose, reach-
ing well beyond apices of endopod and ex-
opod; pleopods 3-5 lacking appendices in-
ternae.
Uropod: Outer ramus with transverse
dentate suture.
Telson: With non-articulating dorsal
spines; single posterolateral articulating
spme.
Species.-
Lophaxius longipes (Bouvier, 1905), offBar-
bados, 225 m.
Lophaxius rudis (Rathbun, 1906), off Ha-
waii, 73-426 m.
Sakaiocaris, new genus
Type species. -By present designation,
Axiopsis brucei Sakai, 1986.
Etymology. - The generic name is a com-
bination of 'sakai,' for Dr. Katsushi Sakai,
plus the Greek 'karis' -a shrimp. Gender:
feminine.
Diagnosis. - Males, females, and her-
maphroditic forms occurring in same
species. Carapace: cervical groove present;
post-cervical carina a low rounded ridge
lacking spines or tubercles; rostral margins
dentate; median carina dentate; submedian
carina dentate; lateral carina dentate. Eye
with reduced pigmentation; eyestalk round-
ed. Antennal acicle a well developed spike.
Maxillipeds: 1-3 with epipod and exo-
pod, 2-3 with reduced podobranch; two ar-
throbranchs on 3.
Pereopods: exopods lacking; epipod plus
podobranch on 1-3; epipod only on 4; two
arthrobranchs on 1-4; one pleurobranch on
2-4. Pereopod 1, chelae asymmetrical. Pe-
reopod 2 chelate. Pereopods 3-5, dactyli
simple.
Pleopods: Pleopod 1of male of 2 articles,
distal article spatulate, with proximo mesial
clump of small hooks. Pleopod 1 in female
(and in only hermaphrodites seen), slender,
elongate, of 2 articles, distal article bearing
marginal setae; pleopod 2 in male with large
exopod and endopod, setose appendix mas-
culina and appendix interna both articulat-
ing proximally on endopod. Pleopods 3-5
with free appendix interna.
Uropod: outer ramus with dentate trans-
verse suture.
Telson: with non-articulating dorsal
spines; with two articulating postero-lateral
spines.
~n~? ~ ~~W~
91 ot 01 91 cr, d2 9' Ot 02 1 '! 2
LophaJCius
Fig. I. Hypothetical scheme for derivation of the Calocarididae from more generalized Axiidae, illustrating
pleopods I and 2, and changes in character-states. Endopod of pleopod 2 shaded. (Abbreviations: app. int.-
appendix interna; app. mas.-appendix masculina; pl.-pleopod.)
Species.-
Sakaiocaris brucei (Sakai, 1986); off West-
ern Australia, in hexactinellid sponges,
296-458 m.
Three synapomorphies separate the Cal-
ocarididae from the Axiidae (s.1.). I. Invar-
iable hermaphroditism. 2. Enlargement of
the appendix masculina of pieopod 2, along
with loss of the setose distal element of the
endopod. 3. Eye reduction and loss of eye
pigment.
A possible pathway in the development
of hermaphroditism, from purely gono-
choristic forms (e.g., Coralaxius), through
forms having some hermaphrodites in the
population (e.g., Posthonocaris, Sakaiocar-
is), to purely hermaphroditic forms (Calo-
caris, Calastacus, Callistocaris, Lophaxius),
is illustrated in Fig. I. Coralaxius Kensley
& Gore, 1981, is purely gonochoristic, and
possesses sexually dimorphic first pleopods.
In the male, the first pleopod is uniramous
and biarticulate, the distal article being
spatulate and having a clump of mesial
hooks. These latter are presumed to come
from the appendix interna which has fused
with the endopod; the exopod has either
been lost or has fused with the endopod. It
is postulated that the genera Posthonocaris
and Sakaiocaris have hermaphroditic forms
in the population. In these, the first pleopod
of the female or protandrous hermaphrodite
is a slender setose, uniramous, biarticulate
structure, while in the males, a spatulate first
pleopod very similar to those found in the
Calocarididae is seen. The presence of pro-
tandrous hermaphrodites in populations of
decapod species has been well documented.
Policansky (1982) mentions a variety of
decapod crustaceans in which protandry oc-
curs, including Calocaris macandreae in
which the biology, and especially reproduc-
tion, has been well examined (see Wolle-
baek 1909, Runnstrom 1925, Buchanan
1963). Bauer (1986) reported the presence
of primary males, primary females, and pro-
tandric hermaphrodites that pass through a
male phase, a transitional phase, and then
become breeding females, in the hippolytid
caridean shrimp Thor manningi. While ten
specimens of Sakaiocaris brucei and four
specimens of the two species of Posthono-
caris form too small a sample on which to
state with certainty that protandrous her-
maphrodites occur in these populations, the
dimensions of the specimens (at least ofthe
former species) would seem to suggest this.
The only ovigerous female of S. brucei seen
has a carapace length of21.9 mm; the three
hermaphrodites with their female first pleo-
pods measure 16.1, 19.6, and 20.0 mm (sug-
gesting that these are either in the transi-
tional phase or approaching the breeding
female phase), while five males range from
13.8-23.3 mm.
Posthonocaris seems to represent a more
advanced stage than Sakaiocaris, in this
supposed trend towards hermaphroditism.
This is seen in the reduction of the setose
distal portion of endopod of pleopod 2, an
elongation of the proximal non-setose por-
tion, and marked enlargement of the ap-
pendix masculina. In the fully hermaphro-
ditic calocaridids Calastacus and
Cal!istocaris, the final stage of this trend,
with complete loss of the distal endopod,
and enlargement and specialization of the
appendix masculina, with which the appen-
dix intern a has fused, can be seen.
There would seem to be a correlation be-
tween development of hermaphroditism and
depth distribution in this group of thalas-
sinideans. Coralaxius, with its spatulate first
pleopod in the male, is a shallow (11-76 m)
coral reef inhabitant. Sakaiocaris inhabits
hexactinellid sponges in 296-456 m. Per-
haps there is a reproductive advantage in
having a hermaphroditic phase in this
species, given its cryptic habit. Species of
Posthonocaris have been recorded in depths
of 73-426 m; Calocaris from 89-1072 m;
Callistocaris from 809-1000 m; Calastacus
from 970-1208 m; Lophaxius from 1733
m.
Loss of eye pigment and corneal facets,
along with a breakdown of the distinction
between cornea and stalk, can be seen in the
four genera ofthe Calocarididae, suggesting
loss of function linked to increased depth
distribution. The hypothetical scheme pro-
posed in Fig. 1 requires that anteriorly flat-
tened eyes arose independently in Calocaris
and Callistocaris, which is not unfeasible.
Similar flattening of eyes can be seen in sev-
eral decapod groups such as cave-dwelling
hippolytid shrimps, deepsea bresiliid
shrimp, as well as in the stomatopod genus
Bathysquilla (R. B. Manning, pers. comm.).
Acknowledgments
I am very grateful to the following for the
loan of material: A. J. Bruce of the Northern
Territory Museum of Arts and Sciences,
Australia; A. Johnson of the museum of
Comparative Zoology, Harvard University;
R. Springthorpe ofthe Australian Museum,
Sydney; M. van der Merwe of the South
African Museum, Cape Town. I thank F. A.
Chace, Jr., A. B. Williams, and R. B. Man-
ning, all of the Smithsonian Institution, for
their (as always) perceptive comments and
improvements to the manuscript.
Alcock, A. 190I. A descriptive catalogue of the In-
dian deep-sea Crustacea Decapoda Macrura and
Anomala, in the Indian Museum. Being a re-
vised account of the deep-sea species collected
by the Royal Indian Marine Survey Ship In-
vestigator. Calcutta, Trustees of the Indian Mu-
seum. 286 pp, 3 pis.
--, & A. R. S. Anderson. 1896. Illustrations of
the zoology of the Royal Indian Marine Survey
Ship Investigator under the command of Com-
mander T. H. Heming, R.N. Crustacea, Part 4,
plates 16-27; Calcutta. Office of the Superin-
tendent of Government Printing, India.
Anderson, A. R. S. 1896. Natural history notes from
the R.I.M. Survey Steamer 'Investigator,' Com-
mander C. F. Oldham, R.N., commanding. Se-
ries II, No. 21. An account of the deep sea Crus-
tacea collected during the season 1894-95.-
Journal of the Asiatic Society of Bengal 65(2):
88-106.
Balss, H. 1957. Decapoda.-Dr. H. G. Bronns Klas-
sen und Ordnungen des Tierreichs 5(I)7( 12):
1505-1672.
Bauer, R. T. 1986. Sex change and life history pattern
in the shrimp Thor manningi (Decapoda: Car-
idea): A novel case of partial protandric her-
maphroditism. - Biological Bulletin 170(1):11-
31.
Bell, T. 1853. A history of the British stalk-eyed crus-
tacea. John Van Voorst, London, lxv, 386 pp,
174 figs.
Borradaile, L. A. 1903. On the classification of the
Thalassinidea.-Annals and Magazine of Nat-
ural History (7) 12:534-551.
Bouvier, E. L. 1905. Sur les Thalassinides recueillis
par Ie Blake dans la mer des Antilles et Ie golfe
du Mexique.-Comptes Rendus Hebdomadaire
des Seances de I'Academie des Sciences 141(21):
802-806.
Buchanan, J. B. 1963. The biology of Calocaris ma-
candreae (Crustacea: Thalassinidea). - Journal
of the Marine Biological Association of the
United Kingdom 43(3):729-747.
Faxon, W. 1893. Reports on the dredging operations
off the west coast of Central America to the
Galapagos, to the west coast of Mexico, and in
the Gulf of California, in charge of Alexander
Agassiz, carried on by the U.S. Fish Commis-
sion Steamer "Albatross," during 1891, Lieut.
Commander Z. L. Tanner, U.S.N., command-
ing.-Bulletin of the Museum of Comparative
Zoology at Harvard College 24(7): 149-220.
Grebenyuk, L. P. 1975. Two new species of the su-
perfamily Thalassinidea. - Zoologichesckii
Zhurnal 54(2):299-304 [In Russian].
Huxley, T. H. 1878. On the classification and the
distribution of the crayfishes.-Proceedings of
the Zoological Society of London for the year
1878:752-788.
Kensley, B., & R. H. Gore. 1981. Coralaxius abelei,
new genus and new species (Crustacea: Deca-
poda: Thala3sinidea: Axiidae): A coral-inhab-
iting shrimp from the Florida Keys and the west-
ern Caribbean Sea. - Proceedings of the
Biological Society of Washington 93(4): 1277-
1294.
Man, J. G. de. 1925. The Decapoda of the Siboga-
Expedition. Part 6. The Axiidae collected by the
Siboga-Expedition. -Siboga-Expeditie Mono-
graphie 39a5:1-127.
McArdle, A. F. 1900. Natural history notes from the
Royal Indian Marine Survey Ship 'Investigator,'
Commander T. H. Heming, R.N., command-
ing. Series III, No.4. Some results of the dredg-
ing season 1899-1900.-Annals and Magazine
of Natural History (7)6:471-478.
Ortmann, A. 1891. Die Decapoden-Krebse des
Strassburger Museums. 3. Die Abtheilungen der
Reptantia Boas: Homaridae, Loricata und Thal-
assinidea.-Zoologischen Jahrbiichern 6: I-58.
Policansky, D. 1982. Sex change in plants and ani-
mals.-Annual Review of Ecology and System-
atics 13:471-495.
Rathbun, M. J. 1904. Decapod crustaceans of the
northwest coast of North America. - Harriman
Alaska Expedition 10:1-190.
---. 1906. The Brachyura and Macrura of the
Hawaiian Islands.-Bulletin of the U.S. Fish
Commission for 1903, part 3:829-930.
Runnstr6m, S. 1925. Beitrag zur Kenntnis einiger
hermaphroditischen dekapoden Crustaceen.-
Bergens Museums Skrifter, n.s., 3(2):1-115.
Saint Laurent, M. de. 1972. Un thalassinide nouveau
du golfe de Gascogne, Calastacus laevis sp. novo
Remarques sur Iegenre Calastacus Faxon (Crus-
tacea Decapoda Axiidae).-Bulletin du Mu-
seum National d'Histoire Naturelle (3)35, Zool-
ogie 29:347-356.
Sakai, K. 1986. Axiopsis brucei sp. nov., a new sponge-
inhabiting axiid (Crustacea: Decapoda: Thal-
assinidea), from north-west Australia. - The
Beagle. Occasional Papers of the Northern Ter-
ritory Museum of Arts and Sciences 3(1):11-20.
Schmitt, W. L. 1921. The marine decapod crustacea
of California. - University of California Publi-
cations in Zoology 23: 1-359.
Squires, H. J. 1965. A new species ofCalocaris(Crus-
tacea: Decapoda, Thalassinidea) from the
northwest Atlantic.-Journal of the Fisheries
Research Board of Canada 22(1):1-11.
Stebbing, T. R. R. 1914. South African Crustacea
(Part 7 of S. A. Crustacea, for the marine in-
vestigations in South Africa).-Annals of the
South African Museum 15(1):I-55.
Stephensen, K. 1910. Storkrebs. 1. Skjoldkrebs.
[Danmarks Fauna 9]. G. E. C. Gads, Copen-
hagen, 193 pp., 108 figs.
Wollebaek, A. 1909. Effektiv hermaphroditisme hos
en decapod Crustace, Calocaris Macandreae,
Bell.- Nyt Magazin for Naturvidenskaberne,
Christiania 47(3):251-268.
Department ofInvertebrate Zoology, Na-
tional Museum of Natural History, NHB-
163, Smithsonian Institution, Washington,
D.C. 20560.