Shrimps of the Pasiphaeid Genus

Leptochela with Descriptions of

Three New Species

(Crustacea: Decapoda: Caridea)

FENNER A. CHACE, JR.

fe

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 222
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 22 2

Shrimps of the Pasiphaeid Genus

Leptochela with Descriptions of

Three New Species

(Crustacea: Decapoda: Caridea)

Fenner A. Chace, Jr.

APR g 2 1378

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976
ABSTRAC T

Chace, Fenner A., Jr. Shrimps of the Pasiphaeid Genus Leptochela with

Descriptions of Three New Species (Crustacea: Decapoda: Caridea). Smithsonian

Contributions to Zoology, number 222, 51 pages, 37 figures, 1976.—Twelve

species of Leptochela are described and illustrated, including L. hawaiiensis,

new species, from Hawaii; L. irrobusta, new species, from the Red Sea to the

Marshall and Samoa islands; and L. papulata, new species, from the Carolinas,

Georgia, and the eastern Gulf of Mexico. Proboloura, new subgenus, is proposed

for the types-species L. carinata Ortmann. The geographic distribution and

sexual dimorphism of the species are discussed.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 6168. SERIES COVER DESIGN:

The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Chace, Fenner Albert.

Shrimps of the Pasiphaeid genus Leptochela with descriptions of three new species (Crustacea,

Decapoda, Caridea)

(Smithsonian contributions to zoology ; no. 222)

Bibliography: p.

Supt. of Docs, no.: SI 1.27:222

1. Leptochela. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to

zoology ; no. 222.

QL1.S54. no. 222 [QL444.M33] 591'.08s [595'.3843] 75-619368
Contents

Page

Introduction 1

Zoogeography 1

Systematic Discussion 1

Acknowledgments 2

Genus Leptochela Stimpson ' 2

Key to the Species 3

Subgenus Leptochela Stimpson 4

1. L. (L.) aculeocaudata Paulson 4

2. L. (L.) bermudensis Gurney 7

3. L. (L.) gracilis Stimpson 11

4. L. (L.) hawaiiensis, new species 15

5. L. (L) irrobusta, new species 19

6. L. (L.) japonica Hayashi and Miyake 23

7. L. (L.) papulata, new species 26

8. L. (L.) pugnax De Man 31

9. L. (L.) robusta Stimpson 34

10. L. (L.) serratorbita Bate 36

ILL . (L.) sydniensis Dakin and Colefax 40

Proboloura, new subgenus 44

12. L. (P.) carinata Ortmann 45

Literature Cited 49

i n

Shrimps of the Pasiphaeid Genus

Leptochela with Descriptions of

Three New Species

(Crustacea: Decapoda: Caridea)

Fenner A. Chace, Jr.

Introduction

ZOOGEOGRAPHY.—The little shrimps of the genus

Leptochela, ranging in adult size from carapace

lengths of less than three to nearly ten mm, are

often abundant in certain parts of the world, especially near the sea surface at night. Seven of the 12

species recognized in this review are known from

the Indo-West-Pacific region: from the Red Sea

eastward to the Marshall and Samoa islands and

from Japan and Korea southward to South Australia and Tasmania (Figure 1). An eighth species has been found only in Hawaii. The remaining four species occur in the western Atlantic from

the latitude of Massachusetts to northern Brazil

near the mouth of the Amazon River. Two of the

species occurring in the Red Sea apparently have

invaded the eastern Mediterranean, and another

species wide ranging in the western Atlantic has

been taken in the eastern Pacific near the southern tip of Baja California. It seems apparent that

Leptochela has a disjunct distribution. Except for

three possible migrants through man-made canals,

the genus is unknown until now from the west

coasts of North, Central, and South America; most

of the east coast of South America; and the coasts

of Europe and most of Africa. More intensive colFenner A. Chace, Jr., Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

lecting in some of these areas will almost certainly

expand the known range of Leptochela, but it is

unlikely that the genus occurs naturally in the

eastern Atlantic, where advanced collecting techniques have been widely used for a long time.

SYSTEMATIC DISCUSSION.—The species of Leptochela are so consistently distinct from the other

genera of the Pasiphaeidae that they have sometimes been assigned to a separate family, the Leptochelidae Paulson, 1875. There is little justification for such discrimination, however, for the

mouthparts and pereopods of Leptochela are characteristically pasiphaeid. On the other hand, this

study, the first worldwide survey of the genus, suggested rather convincingly that one of the species

is sufficiently distinct from the others to be accorded subgeneric rank.

Another clarification disclosed by the study is

that the sexual dimorphism first noticed by Kemp

(1925:250) and represented by a longitudinally

tricarinate carapace in breeding females only, occurs in the majority, but not all, of the species of

Leptochela. In two species, the carapace is tricarinate in both sexes, while in two other species

it is not so modified in either sex (see L. sydniensis, Remarks).

Thanks to the rather extensive series of Leptochela in the Smithsonian collections and to the

kind cooperation of colleagues mentioned below,

I have been able to examine specimens of all of

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE 1.—World distribution of the genus Leptochela as suggested by available records.

the described species except L. hainanensis Yu,

1936. I believed at first that this name was available for the Indo-Pacific species generally misidentified as L. aculeocaudata, but certain of the

characters ascribed to L. hainanensis disagree with

those in material that I have seen of the misidentified species. Only the comparison of specimens of

L. hainanensis with those described below under

the name L. sydniensis can decide whether these

species are distinct or whether L. hainanensis

should take precedence as a senior synonym of L.

sydniensis.

The postorbital carapace lengths of the specimens listed in the "Material" section of each of the

species descriptions are shown in parentheses. The

indicated magnifications of the illustrations are

reasonably but not always precisely accurate.

ACKNOWLEDGMENTS.—This study was initially

motivated and continuously nurtured by John C.

Yaldwyn of the National Museum of New Zealand;

his profound knowledge of the other pasiphaeid

genera was especially valuable during the entire

progress of the investigation. Also, I take this opportunity to thank Keiji Baba of the Kumamoto

University Faculty of Education for translating

pertinent Japanese literature; Harold S. Feinberg

of the American Museum of Natural History for

arranging the loan of study material; Takahiro

Fujino of the Kyushu University Faculty of Medicine for translating Japanese literature and for

sending comparative material of Leptochela sydniensis on loan; D. J. G. Griffin of the Australian

Museum for the loan of all Leptochela specimens

in that institution; L. B. Holthuis of the Rijksmuseum van Natuurlijke Historie, Leiden, for permitting the examination of more than 1800 specimens from the Red Sea; Richard L. Wigley of the

National Marine Fisheries Service, Woods Hole,

for checking the documentation of a unique specimen; Austin B. Williams of the Systematics Laboratory of the National Marine Fisheries Service,

Washington, D.C., for making available material

in the Institute of Marine Science, Morehead City,

North Carolina; and my Smithsonian colleagues

Horton H. Hobbs, Jr., and Raymond B. Manning

for reviewing the manuscript.

Genus Leptochela Stimpson

Leptochela Stimpson, 1860:42.

DEFINITION.—Pasiphaeid with conventional rostrum. Carapace and rostrum unarmed dorsally.

Branchiostegal tooth and branchiostegal sinus absent. Sixth abdominal somite with transverse cari-
NUMBER 222

nate ridge near anterior end of dorsal surface and

long fixed posteriorly directed spine near posterior

end of ventrolateral margin. Telson with mesial

pair of movable spines anteriorly, 1 or 2 pairs of

dorsolateral movable spines, and 5 pairs of prominent posterior movable spines, all but lateral pair

(perhaps actually belonging to dorsolateral series)

of latter minutely serrate on one or both lateral

and mesial margins (minute additional pair of

spines sometimes present between bases of median

pair). Mandibular palp broad, flattened, and undivided. Third maxilliped with 2 arthrobranchs.

Fourth pereopod shorter than 3rd, longer than

5th. Exopods of pleopods not unusually long. Both

branches of uropod with series of movable lateral

spines.

TYPE-SPECIES (selected by Kemp, 1915:310).—

Leptochela gracilis Stimpson.

Key to the Species

I. Sixth abdominal somite bearing movable lappet near anterior end of dorsal surface; telson

with anterior pair of dorsolateral spines nearly in line with anteriorly placed mesial

pair; antenna! scale usually at least y4 as long as carapace; 3rd pereopod with exopod

reaching nearly or quite to distal end of ischium; 5th pereopod noticeably reduced,

about 2/3 as long as 4th. Subgenus Proboloura. One species 12. L. (P.) carinata

Sixth abdominal somite without dorsal lappet; telson with anterior pair of dorsolateral

spines arising considerably posterior to mesial pair; antennal scale usually less than %

as long as carapace; 3rd pereopod with exopod not nearly reaching distal end of

ischium; 5th pereopod at least 34 as long as 4th. Subgenus Leptochela 2

2.(1) Telson armed with 1 pair of dorsomesial and 2 pairs of dorsolateral spines in addition

to 5 pairs of prominent spines in posterior series; orbit armed with mesially directed

tooth on ventral margin or with tooth at suborbital angle 3

Telson armed with 1 pair of dorsomesial and 1 pair of dorsolateral spines in addition to

posterior series; ventral margin of orbit and suborbital angle unarmed (except in L.

pugnax) 5

3.(2) Suborbital angle dentate; orbital margin serrate dorsolaterally; telson usually with pair

of minute spines between bases of median pair of large posterior spines

10. L. (L.) serratorbita

Suborbital angle rounded, unarmed; orbital margin usually entire dorsolaterally; telson

without pair of minute spines between bases of median pair of large posterior spines . 4

4.(3) Appendix masculina (not including spines) on second male pleopod rarely overreaching

appendix interna; size small, ovigerous females with postorbital carapace length less

than 5 mm 5. L. (L.) irrobmta

Appendix masculina (not including spines) far overreaching appendix interna; size large,

ovigerous females with postorbital carapace length more than 6 mm .9 . L. (L.) robusta

5.(2) Suborbital angle dentate 8. L. (L.) pugnax

Suborbital angle rounded 6

6.(5) Dorsal margin of 5th abdominal somite slightly or prominently uneven in lateral view .7

Dorsal margin of 5th abdominal somite regularly convex or nearly straight in lateral

view 8

7.(6) Fifth abdominal somite with dorsal margin very uneven in lateral view; telson without

pair of minute spines between bases of median pair of large posterior spines

6. L. (t.) japonica

Fifth abdominal somite with 1 to 3 low prominences on dorsal margin; telson with pair

of minute spines between bases of median pair of large posterior spines

7. L. (L.) populate

8.(6) Fifth abdominal somite armed with prominent sharp posteromedian tooth; size large,

ovigerous females with postorbital carapace length more thaQ 7 mm .3 . L. (L.) gracilis

Fifth abdominal somite without posteromedian tooth; size small, ovigerous females with

postorbital carapace length less than 5 mm 9

9.(8) Telson with pair (sometimes fused) of minute spines between bases of median pair of

large posterior spines 10

Telson without pair of minute spines between bases of median pair of large posterior

spines 11
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

10.(9) Dorsal margin of orbit entire; endopod of 1st male pleopod with distolateral angle produced; appendix masculina (not including spines) far overreaching appendix interna

2. L. (L.) bermudensis

Dorsal margin of orbit finely serrate under high magnification; endopod of 1st male

pleopod subsymmetrically rounded distally; appendix masculina (not including spines)

not overreaching appendix interna 4. L. (L.) hawaiiensis

11.(9) Rostrum usually rather broadly lanceolate in lateral view; carapace with 3 longitudinal

dorsal ridges in both males and females; basal antennal segments concealed by carapace;

movable finger of 1st pereopod bearing 14-22 spines on opposable margin, that of 2nd

pereopod with 17-26 spines; ovigerous females with postorbital carapace length 2.4-2.6

m m 1. L. (L.) aculeocaudata

Rostrum narrower in lateral view, dorsal margin usually sinuous in lateral view; carapace

with dorsolateral longitudinal ridges only in breeding females; basal antennal segments

not concealed by carapace; movable finger of first pereopod bearing 20-44 spines on

opposable margin, that of second pereopod with 21-46 spines; ovigerous females with

postorbital carapace length 2.6-4.4 mm 11. L. (L.) sydnicnsis

Subgenus Leptochela Stimpson

DEFINITION.—Sixth abdominal somite without

dorsal lappet. Telson with origins of anterior pair

of dorsolateral spines and of anterior dorsal pair

separated by distance amounting to at least */5

length of telson, not including posterior spines.

Antennal scale usually less than 2/3 as long as carapace. Third pereopod with exopod not reaching

nearly so far as distal end of ischium. Fifth pair of

pereopods at least y4 as long as 4th pair.

1. Leptochela (Leptochela) aculeocaudata Paulson

FIGURES 2-4

Leptochela aculeocaudata Paulson, 1875:100, pi. 16: figs. 1-ls

[translation, 1961:106, pi. 15: figs. 1-ls].—Nobili, 1906:28,

fig. 4.—Balss, 1936:4, 57, 64, fig. 3.—Gurney, 1939:428, 429,

433.

Leptochela aculeocaudata hainanensis.—Armstrong, 1941:1.

DIAGNOSIS.—Rostrum lanceolate, dorsal margin

convex posteriorly, straight or slightly concave anteriorly. Carapace with 3 longitudinal dorsal ridges

in both males and females. Orbital margin entire,

not serrate, without mesially directed tooth on ventral portion; suborbital angle unarmed. Fifth abdominal somite entire, without dorsal elevations or

posterior tooth. Telson with 1 pair of dorsolateral

spines in addition to anterior mesial pair; posterior

margin without pair of minute mesial spines in

addition to usual 5 pairs of prominent spines. Antennal scale less than i/2 as long as carapace. First

pereopod with 16 to 23 spines on opposable margin of movable finger. Second pereopod with 17 to

26 spines on opposable margin of movable finger.

Third pereopod with exopod not nearly reaching

distal end of ischium. Endopod of 1st pleopod of

male obliquely rounded distally, lateral margin

slightly concave but not flared distally. Appendix

masculina, excluding spines, usually not overreaching appendix interna. Maximum carapace length

2.9 mm.

DESCRIPTION.—Rostrum (Figure 2a-c) lanceolate, rather high proximally, dorsal margin straight

or slightly concave anteriorly, usually overreaching

basal segment of antennular peduncle. Carapace

with median carina extending posteriorly barely to

midlength and paired blunt dorsolateral ridges extending to posterior i/4 in males and nonbreeding

females, both median carina and dorsolateral

ridges extending at least to posterior % of carapace length in breeding females; dorsolateral

ridges elevated in both sexes, characteristically

concealing all of dorsal midline except rostral crest

from lateral view; carapace usually completely

concealing basal segment of antennal peduncle

from lateral view. Orbital margin (Figure 2b,c)

entire, not spinulose, without mesially directed

tooth on ventral portion; suborbital angle broadly

rounded, unarmed.

Abdominal terga (Figure 2d) usually rounded

on all somites, sometimes with suggestion of blunt

carina on 5th somite. Fifth somite entire, without

dorsal prominences or posterodorsal tooth. Sixth

somite nearly twice as long as high, with low transverse carinate swelling at extreme anterior end

usually concealed beneath posterior margin of 5th

somite; rather short, slightly curved spine on ventrolateral surface and distinct acute tooth on pos-
NUMBER 222

FIGURE 2.—Leptochela (Leptochela) aculeocaudata, ovigerous female from Red Sea off Sinai

Peninsula, carapace length 2.4 mm: a, carapace and anterior appendages; b, anterior part of

carapace and rostrum, lateral aspect; c, anterior part of carapace and eyes, dorsal aspect;

d, abdomen; e, posterior end of 6th abdominal somite; /, telson and uropods; g, posterior margin

of telson; h, right antennule, dorsal aspect; i, right antenna, ventral aspect. (Magnifications:

a, d, X 12; b, c, e, f, h, i, X 25; g, X 62.)

terodorsal margin of posterolateral lobe (Figure

2e). Telson (Figure 2/) nearly li/£ times as long

as 6th somite, about 3 times as long as wide, armed

with 1 pair of dorsolateral spines at about midlength, in addition to anterior mesial pair; posterior margin (Figure 2g) without pair of minute

spines between bases of mesialmost of usual 5 pairs

of prominent spines.

Eye (Figure 2c) with indistinct papilla on mesial surface of stalk near juncture with cornea,

cornea little swollen, barely wider than stalk.

Antennular peduncle (Figure 2h) with stylocerite reaching as far as distolateral margin of basal

segment; 2nd segment about as long as distal segment in ventromesial aspect, much shorter in dorsal aspect.

Antennal scale (Figure 2i) less than i/2 as long

as carapace, 3.0 to 4.0 times as long as wide, lateral

margin distinctly concave near midlength, blade

forming oblique shoulder at base of distal tooth.

Distal segments of antennal peduncle less than 2/3

as wide as scale, extending distinctly beyond midlength of scale; basal segment bluntly produced

distoventrally, without acute tooth.

Mouthparts as illustrated (Figure 3a-/). Third

maxilliped (Figure 3/) usually not overreaching

antennal scale, distal segment nearly y4 as long as

penultimate segment.

First pereopod (Figure 3g) occasionally overreaching antennal scale by more than length of

fingers; fingers 1.8 to 2.0 times as long as palm;

dactyl (Figure Sh) armed with 16 to 23 spines on

opposable margin. Second pereopod (Figure Si)

overreaching antennal scale by at least ys length

of fingers; fingers 2.0 to 2.4 times as long as palm;

dactyl (Figure 3/) armed with 17 to 26 spines on
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Leptochela (Leptochela) aculeocaudata, ovigerous female from Red Sea off Sinai

Peninsula, carapace length 2.4 mm: a, right mandible; b, right 1st maxilla; c, right 2nd maxilla;

d, right 1st maxilliped; e, right 2nd maxilliped; /, right 3rd maxilliped; g, right 1st pereopod;

h, same, fingers; i, right 2nd pereopod; j , same, fingers; k, right 3rd pereopod; I, right 4th

pereopod; m, right 5th pereopod. (Magnifications: /, g, i, k-m, X 25; a-e, h, j , X 62.)

opposable margin. Third pereopod (Figure 3k)

overreaching extreme anterior margin of carapace

by length of dactyl and about y2 of propodus;

exopod not nearly reaching distal end of ischium;

ischium bearing 2 spines near flexor margin; merus

with 3 spines; carpus with 1 spine; dactyl much

shorter than propodus. Fourth pereopod (Figure

3/) reaching to about distal end of ischium of 2nd

pereopod when both extended anteriorly; dactyl

longer than propodus. Fifth pereopod (Figure

3m) similar to and little shorter than 4th, reaching about midlength of ischium of 2nd pereopod

when both extended anteriorly; dactyl fully as long

as propodus.

Endopod of 1st pleopod of male (Figure 4a,b)

obliquely rounded distally, lateral margin slightly
NUMBER 222

concave but not flared distally. Appendix masculina (Figure 4c,d) bearing 6 long spines and, excluding latter, usually not overreaching appendix

interna. Lateral branch of uropod (Figure 2/)

armed with 10 to 14 movable spines.

SIZE.—Carapace lengths of males, 2.0-2.9 mm;

of females without eggs, 2.0-2.6 mm; of ovigerous

females, 2.4, 2.6 mm; of juvenile, 1.4 mm.

MATERIAL.—RED SEA. Sinai Peninsula: 1 $ (2.8) 1 ovig.

$ (2.4) (Rijksmuseum van Natuurlijke Historie, Leiden).

QUEENSLAND, AUSTRALIA. Bustard Bay (24°05'S,

151°48'E); 19 September 1938; Haul 46/38 (old number),

Net 200; K. Sheard: 1 3 (2.9) 2$ (2.1+, 2.6) (1 ovig. (2.6))

(Australian Museum).

SAMOA ISLANDS. Apia Harbour, Upolu; 12 October

1936: submerged light; Second Templeton Crocker-American

Museum Expedition: 24 $ (2.0-2.6) 2$ (2.4, 2.6) 1 juv. (1.4)

(American Museum of Natural History).

HABITAT.—Balss (1936:5) recorded L. (L.)

aculeocaudata from off Alexandria, Egypt, in

depths of 51 to 88 meters on muddy bottoms with

and without vegetation. The Samoan collection

was made at the surface over a submerged light.

TYPE-LOCALITY.—Red Sea.

DISTRIBUTION.—Known with reasonable certainty

only from the Red Sea and the two regions recorded here: Queensland, Australia, and the Samoa Islands. Also, there is little doubt that the

specimens recorded by Balss (1936:5) from off

Alexandria, Egypt, in 51 to 88 meters were correctly identified as belonging to this species.

REMARKS.—Although only one pair—a male and

an ovigerous female—of the species described above

was found in the more than 1800 specimens of

Leptochela kindly made available by L. B. Holthuis in advance of the preparation of his report

on Red Sea collections in his custody, I have little

hesitation in identifying this pair as L. (L.) aculeocaudata. The elevated dorsolateral ridges that

conceal the midline of the carapace, except for

the rostral crest, from lateral view (well illustrated

in Paulson's Figure la) and the small size of

adult specimens seem to characterize the species

sufficiently.

The specimens from the Samoa Islands identified as L. aculeocaudata hainanensis by Armstrong

(1941) and made available by H. S. Feinberg of

the American Museum of Natural History are so

poorly preserved that positive identification is difficult, but the presence of highly elevated dorsod

FIGURE 4.—Leptochela (Leptochela) aculeocaudata, male from

Red Sea off Sinai Peninsula, carapace length 2.8 mm: a,

right 1st pleopod; b, same, endopod; c, right 2nd pleopod;

d, same, appendix masculina and appendix interna. (Magnifications: a, c, X 12; b, d, X 62.)

lateral ridges on the carapace in both males and

females seems to establish their identity with this

species. The only noticeable difference between

these specimens and those from the Red Sea and

Queensland is that the basal segment of the antenna is not concealed by the carapace in the

Samoan material, but I am inclined to believe that

this is an artifact resulting from the skeletonized

condition of the latter specimens, in which all of

the appendages are very loosely articulated.

See L. (L.) sydniensis for discussion of material

that I believe to have been misidentified with L.

(L.) aculeocaudata.

2. Leptochela (Leptochela) bermudensis Gurney

FIGURES 5-7

TLeptochela carinata Ortmann, 1893:41 [part, not pi. 4: fig. 1].

Leptochela carinata.—Schmitt, 1924:69.—Gurney, 1936:786,

pi. 1: figs. 1-5, pi. 2: figs. 7-12, pi. 3: figs. 14-18, pi. 4:

figs. 19-30, pi. 5: figs. 33-34. [Not L. carinata Ortmann,

1893, pi. 4: fig. 1.]

•Leptochela bermudensis Gurney, 1939:427, figs. 1-10.—Chace,

1940:131, fig. 10—Holthuis, 1955: fig. 15b.—Springer and

Bullis, 1956:10.—Chace, 1972:16.

DIAGNOSIS.—Rostrum with dorsal margin usually

straight or concave, occasionally sinuous or even

convex throughout. Carapace with 3 longitudinal

dorsal ridges in breeding females only. Orbital

margin entire, not serrate, without mesially di-
8

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rected tooth on ventral portion; suborbital angle

rounded, rarely with minute denticle. Fifth abdominal somite entire, without dorsal elevations or posterior tooth. Telson with 1 pair of dorsolateral

spines in addition to anterior mesial pair; posterior margin with pair (rarely fused) of minute

mesial spines in addition to usual 5 pairs of prominent spines. Antennal scale little more than 1/% as

long as carapace. First pereopod with 11 to 30

spines on opposable margin of movable finger. Second pereopod with 18 to 37 spines on opposable

margin of movable finger. Third pereopod with

exopod not nearly reaching distal end of ischium.

Endopod of 1st pleopod of male flared distolaterally. Appendix masculina, not including spines,

distinctly overreaching appendix interna. Maximum carapace length 4.5 mm.

DESCRIPTION.—Rostrum (Figure 5a) with dorsal

margin somewhat variable, usually nearly straight

or concave, occasionally sinuous or even slightly

convex from base to tip, rarely overreaching basal

segment of antennular peduncle. Carapace with

median dorsal carina on anterior 14 to i/£ of length

but without paired dorsolateral carinae in males

and nonbreeding females, dorsally tricarinate over

most of length in breeding females. Orbital margin (Figure 5b) entire, not serrate, ventral portion without mesially directed tooth; suborbital

angle rounded, rarely with minute marginal

denticle.

FICURE b.—Leptochela (Leptochela) bermudensis, ovigerous female from Barbuda, Leeward

Islands, carapace length 3.0 mm: a, carapace and anterior appendages; b, anterior part of carapace and eyes, dorsal aspect; c, abdomen; d, posterior end of 6th abdominal somite: e, telson

and uropods; /, posterior margin of telson; g, same, mesial spines; h, right antennule, dorsal

aspect; i, right antenna, ventral aspect. (Magnifications: a, c, e, X 12- b, d f h i X 25- e

X 260.) >>!>, , AJ, g ,
NUMBER 222

Abdominal terga (Figure be) regularly rounded

on 3 anterior somites and all but posterior part of

4th. Fifth somite bluntly, sometimes obscurely,

carinate dorsally, entire, without dorsal prominences of any kind and without posterodorsal

tooth. Sixth somite nearly twice as long as high,

with usual transverse swelling and carina near anterior end of dorsal surface, long slender spine on

ventrolateral surface and usually distinct, rarely

obscure, acute tooth (Figure bd) on posterodorsal

margin of posterolateral lobe. Telson (Figure be),

not including posterior spines, about 1.6 times as

long as 6th somite, little more than 3 times as long

as wide, armed with 1 pair of dorsolateral spines

at about midlength, in addition to anterior mesial

pair; posterior margin (Figure 5/) bearing pair

(rarely fused) of minute mesial spines (Figure

5g) between bases of mesial pair of usual 5 pairs

of prominent spines.

Eye (Figure 56) with papilla on mesial surface

of stalk proximal to cornea, cornea little if at all

wider than stalk.

Antennular peduncle (Figure bh) with stylocerite reaching nearly as far as distolateral margin

of basal segment; 2nd segment longer than distal

segment in mesial aspect but distinctly shorter in

dorsal aspect.

Antennal scale (Figure bi) little more than i/2

as long as carapace, 2.8 to 3.7 times as long as wide,

lateral margin slightly sinuous, blade forming

rather distinct shoulder at base of distal tooth. Distal segments of antennal peduncle about 2/3 as

wide as scale, not reaching midlength of scale.

Mouthparts as illustrated (Figure 6a-/). Third

maxilliped (Figure 6/) rarely overreaching antennal scale, distal segment about y4 as long as

penultimate segment.

First pereopod (Figure 6g) overreaching antennal scale by slightly more or slightly less than

length of fingers; fingers 1.2 to 1.7 times as long as

palm; dactyl (Figure 6h) armed with 11 to 30

spines on opposable margin. Second pereopod

(Figure 6?) rarely overreaching antennal scale by

as much as length of fingers; fingers 1.7 to 2.2 times

as long as palm; dactyl (Figure 6/) armed with

18 to 37 spines on opposable margin. Third pereopod (Figure 6k) overreaching extreme anterior

margin of carapace by about i/2 length of dactyl;

exopod falling considerably short of distal end of

ischium; ischium armed with row of about 6 slender spines near extensor margin and 4 subequally

spaced stouter spines on lateral surface; merus with 5

blunt spines on lateral surface; dactyl shorter than

propodus. Fourth pereopod (Figure 6/) reaching

to about midlength of ischium of 2nd pereopod

when both extended anteriorly; dactyl usually

shorter than propodus. Fifth pereopod (Figure

6m) similar to and little shorter than 4th, reaching nearly to midlength of ischium of 2nd pereopod when both extended anteriorly; dactyl shorter

than propodus.

Endopod of 1st pleopod of male (Figure 7a,b)

with lateral margin distinctly flared distally. Appendix masculina (Figure 7c,d) bearing 9 long

spines, far overreaching appendix interna, without including spines. Lateral branch of uropod

(Figure be) armed with 8 to 15 movable spines,

in addition to setae.

SIZE.—Carapace lengths of males, 2.1-3.0 mm;

of nonbreeding females, 2.1-3.0 mm; of nonovigerous breeding female, 2.7 mm; of ovigerous females, 2.4-3.0 mm; of juveniles, 1.6-1.9 mm. In

contrast, Chace (1940) recorded males with a maximum carapace length of 4.5 mm and ovigerous

females of 3.2 mm from off Bermuda.

MATERIAL.—GULF OF MEXICO. Cayos Areas, Bahia de

Campeche; 18 m; 11 December 1952; from bonefish stomach;

Oregon: 1 9 (2.2) 2 juv. (1.6, 1.8). Cayos Areas, Bahfa de

Campeche; 2O°12'N, 9^9^ ; 11 m; 18 July 1957; traplift

net with light; Oregon Sta. 1848: 2 juv. (1.6, 1.9).

PUERTO RICO. Playa de Fajardo; 23 February 1933: 8-

foot circular net under cargo light; Johnson-Smithsonian

Deep-sea Expedition: 1 $ (2.5). Cayo Icacos; 24 February

1933; 8-foot circular net with cargo light and submarine

light; Johnson-Smithsonian Deep-sea Expedition: 9$ (2.1-

2.5) 2 9 (2.1-2.4). Canal de Luis Pefia, Isla de Culebra; 25

February 1933; 8-foot circular net under cargo light; JohnsonSmithsonian Deep-sea Expedition: 3$ (2.3-25). Same; 3-foot

net towed from port launch: 21 $ (2.3-2.6).

VIRGIN ISLANDS. Brewers Bay, Saint Thomas; 1 March

1933; 8-foot circular net under cargo light; JohnsonSmithsonian Deep-sea Expedition: 10 <$ (2.4-2.7) 5 9 (2.6-

3.0).

LEEWARD ISLANDS. Off Oyster Pond Landing, Barbuda;

5 April 1956; 7:00-8:30 P.M.; light over side at anchorage in

5.5 m; Smithsonian-Bredin Caribbean Expeditions Sta. 84-56:

2$ (3.0) 1 ovig. 9 (3.0). Prince Rupert Bay, Dominica: 28

March 1956; 8:45-9:30 P.M.; light over side at anchorage in

7 m; Smithsonian-Bredin Caribbean Expeditions Sta. 64-56:

\$ (2.5).

WINDWARD ISLANDS. West by north of telegraph station, Barbados, little more than i/2 mile off; 55-119 m; rocks

and sand; 5 June 1918; University of Iowa Barbados-Antigua
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 6.—Leptochela (Leptochela) bermudensis, ovigerous female from Barbuda, Leeward

Islands, carapace length 3.0 mm: a, right mandible; b, right 1st maxilla; c, right 2nd maxilla;

d, right 1st maxilliped; e, right 2nd maxilliped; /, right 3rd maxilliped; g, right 1st pereopod;

h, same, fingers; i, right 2nd pereopod; /, same, fingers; k, right 3rd pereopod; /, right 4th

pereopod; m, right 5th pereopod. (Magnifications: c-g, i, k-m, X 25; a, b, h, j , X 62.)
NUMBER 222 11

a

FIGURE 7.—Leptochela (Leptochela) bermudensis, male from

Barbuda, Leeward Islands, carapace length 3.0 mm: a, right

1st pleopod; b, same, endopod; c, right 2nd pleopod; d,

same, appendix masculina and appendix interna. (Magnifications: a, c, X 12; b, c, X 62.)

Expedition Sta. 79: 1 ovig.$ (2.5). Off Lazaretto, Barbados;

37 m; bottom rough; 6 June 1918; University of Iowa

Barbados-Antigua Expedition Sta. 87: 1 ovig. 2 (2.4).

HABITAT.—Although most of the specimens

listed above were taken in relatively shallow

water (5.5-18 m), the type-series (Gurney, 1939)

and material subsequently collected off Bermuda

(Chace, 1940) were taken in the open ocean between the surface and 1200m.

Seven of the 11 lots currently available to me

were obtained at night under a light, yet there is

some evidence that L. (L.) bermudensis may be

less strongly photopositive than is L. (L.) serratorbita, which is sympatric with it over much of

its range. At five stations where a light was used

and both species were collected, L. (L.) serratorbita outnumbered L. (L.) bermudensis 676 to

33. At only one station (Cayo Icacos, Puerto Rico)

was the latter more numerous (11 to 8), while at

two stations (Playa de Fajardo, Puerto Rico, and

Barbuda) L. (L.) serratorbita outnumbered L.

(L.) bermudensis by about 150 to 1. On the other

hand, the lot collected with a townet in Canal de

Luis Pena, Puerto Rico, contained 21 L. (L.) bermudensis and only 4 L. (L.) serratorbita, whereas

a light station at that locality on the same night

produced 27 L. (L.) serratorbita and only 3 L.

(L.) bermudensis. Such evidence must be treated

with caution, however, because of the probability

that swarming occurs in both species.

TYPE-LOCALITY.—Seven miles south of Bermuda

between 1000 meters and the surface.

DISTRIBUTION.—Bermuda to Barbados and southwestern Gulf of Mexico, between 1000 meters or

more and the surface in the open ocean and in

inshore depths as shallow as 5.5 meters.

REMARKS.—Differences and similarities between

this species and L. (L.) papulata are discussed

under the latter species.

It seems apparent from Ortmann's (1893) description of L. carinata that two species were

represented in the type-series collected by the

Plankton-Expedition off Baia de Marajo, Estado

do Para, Brazil, in 50-100 meters. Ortmann's name

should certainly be assigned to the species represented by the largest, figured specimen. The identity of the three smaller specimens without dorsal

teeth on the fifth abdominal somite and without a

lappet on the sixth (which Ortmann believed to

be characters associated with sex or age) is uncertain, but the description is applicable to L. (L.)

bermudensis.

3. Leptochela (Leptochela) gracilis Stimpson

FIGURES 8-10

Leptochela gracilis Stimpson, 1860:42.—Bate, 1888:860, pi.

139: fig. 2.—Balss, 1914:19.—Kemp, 1915: 310—Urita,

1921:216.—Kemp, 1925:251.—Urita, 1926:424.—Yokoya,

1933:13, 212.—Yu, 1936:86, 99—Yokoya, 1939:263—Miyadi,

1940a:7, 8, 11.—Yoshida, 1941:21, pi. 4: fig. 4.—Kubo,

1955:99, figs. 1-3.—Liu, 1955:23, pi. 8: figs. 6-14.—Fujino

and Miyake, 1970:242.

Leptochela pellucida Boone, 1935:105, pis. 26, 27.

DIAGNOSIS.—Rostrum with dorsal margin straight

or sinuous, rarely convex throughout. Carapace

without dorsolateral ridges in either males or females. Orbital margin entire, not serrate, without

mesially directed tooth on ventral portion; suborbital angle unarmed. Fifth abdominal somite entire, without dorsal elevations but with sharp posterior tooth. Telson with 1 pair of dorsolateral

spines in addition to anterior mesial pair; posterior margin without pair of minute mesial spines

in addition to usual 5 pairs of prominent spines.

Antennal scale about % as long as carapace. First

pereopod with 47 to 60 spines on opposable mar-
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8.—Leptochela (Leptochela) gracilis, ovigerous female from Fukien, China, carapace

length 9.6 mm: a, carapace and anterior appendages; b, abdomen; c, posterior end of 6th

abdominal somite; d, telson and uropods; e, posterior margin of telson; /, same, mesial part;

g, right antennule, dorsal aspect; h, right antenna, ventral aspect. (Magnifications: a, b, d, g,

h, X 6; c, X 12; e, X 25; /, X 260.)

gin of movable finger. Second pereopod with 48

to 76 spines on opposable margin of movable finger. Third pereopod with exopod not nearly

reaching distal end of ischium. Endopod of 1st

pleopod of male rounded distally, not flared distolaterally. Appendix masculina, not including

spines, not overreaching appendix interna. Maximum carapace length 9.6 mm.

DESCRIPTION.—Rostrum (Figure 8a) with dorsal

margin straight or sinuous, rarely convex throughout, usually reaching to or beyond level of distal end

of basal segment of antennular peduncle. Carapace,

with median dorsal carina on anterior } 1 0 to 2/3 of

length, not tricarinate in either males or females.

Orbital margin entire, not serrate, without mesially directed tooth on ventral portion; suborbital

angle rounded.

Abdomen (Figure 8&) regularly rounded dorsally

on 3 anterior somites, usually carinate on posterior

part of 4th. Fifth somite carinate, without dorsal

prominences but with sharp median tooth projecting from posterior margin. Sixth somite with

dorsal tranverse swelling near anterior end pronounced and produced slightly posteriorly, posterolateral lobe (Figure 8c) with distinct acute tooth

on posterodorsal margin. Telson (Figure 8d), not

including posterior spines, about 1.4 times as long

as 6th somite, about 3.3 times as long as wide,
NUMBER 222 13

FIGURE 9.—Leptochela (Leptochela) gracilis, ovigerous female from Fukien, China, carapace

length 9.6 mm: a, right mandible; b, right 1st maxilla; c, right 2nd maxilla; d, right 1st maxil->

liped; e, right 2nd maxilliped; /, right 3rd maxilliped; g, right 1st pereopod; h, same, fingers;

i, right 2nd pereopod; /, same, fingers; k, right 3rd pereopod; /, right 4th pereopod; m, right 5th

pereopod. (Magnifications: /, g, i, k-m, X 6; c-e, X 12; a, b, h, j , X 25.)

margins somewhat sinuous anteriorly, armed with

1 pair of dorsolateral spines near midlength, in

addition to anterior mesial pair; posterior margin

(Figure 8e) without pair of minute spines between bases of mesial pair of usual 5 pairs of prominent spines (Figure 8/).

Eye with rather slender papilla on mesial surface of stalk proximal to cornea, cornea slightly

wider than stalk.

Antennular peduncle (Figure 8g) with stylocerite reaching nearly as far as distolateral margin of

basal segment; 2nd segment nearly as long as distal segment in mesial aspect but distinctly shorter

in dorsal aspect in spite of pronounced lobe on

distal margin.

Antennal scale (Figure Sh) 0.6 to 0.7 as long as

carapace, 4.4 to 5.0 times as long as wide; lateral

margin noticeably sinuous, distal tooth continuous with mesial margin of blade, with barely discernible convexity suggesting subdistal shoulder

in latter. Distaf segments of antennal peduncle

slightly more than y2 as wide as scale, reaching to

middle 1/3 of scale; basal segment with distinct

ventral spine.

Mouthparts as illustrated (Figure 9a-/). Third

maxilliped (Figure 9/) not overreaching antennal
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

scale, distal segment about 2.3 times as long as

penultimate segment.

First pereopod (Figure 9g) overreaching antennal scale by no more than length of fingers; fingers 0.9 to 1.3 times as long as palm; dactyl (Figure 9h) armed with 47 to 60 spines on opposable

margin. Second pereopod (Figure 9i) overreaching antennal scale by no more than length of fingers; fingers 1.1 to 1.4 times as long as palm; dactyl

(Figure 9/) armed with 48 to 76 spines on opposable margin. Third pereopod (Figure 9k) overreaching extreme anterior margin of carapace by

about length of dactyl; exopod falling considerably

short of distal end of ischium; ischium with row

of setae but no spines near extensor margin but

with about 5 spines near flexor margin; merus

with row of about 6 similar spines near flexor margin; dactyl shorter than propodus. Fourth pereopod (Figure 9/) reaching nearly to distal end of

ischium of 2nd pereopod when both extended anteriorly; dactyl longer than propodus. Fifth pereopod (Figure 9m) similar to and little shorter than

4th, reaching nearly to midlength of ischium of

2nd pereopod when both extended anteriorly;

dactyl longer than propodus.

Endopod of 1st pleopod of male (Figure 10a,b)

subsymmetrically rounded distally, lateral margin

slightly concave but not flared distally. Appendix

masculina (Figure 10c,d) bearing 7 long spines,

FIGURE 10.—Leptochela (Leptochela) gracilis, male from

Fukien, China, carapace length 8.8 mm: a, right 1st pleopod;

b, same, endopod; c, right 2nd pleopod; d, same, appendix

masculina and appendix interna. (Magnifications: a, c, X 6;

b, d, X 25.)

not overreaching appendix interna, not including

spines. Lateral branch of uropod (Figure Sd)

armed with 8 to 15 movable spines (some not visible in figure).

SIZE.—Carapace lengths of males, 7.1-8.8 mm;

of nonovigerous females, 8.6, 8.8 mm; of ovigerous

females, 8.0-9.6 mm.

MATERIAL.—CHINA. Ch'ingtao, Shantung: 1924; C. T.

Urita: 13 (7.1). Fukien; S. F. Light: 4$ (7.8-8.8) 17$

(8.0-9.6) (15 ovig. (8.0-9.6)).

HABITAT.—Although L. (L.) gracilis is of economic importance in parts of its range and has

been mentioned in the literature more frequently

than any of the other species of the genus, I am

unaware of any discussion of its habits or ecological relationships. This may be due to the fact that

much of the important literature is in Japanese or

Chinese. It seems to have been taken most commonly on or near the bottom in depths of about

30 to 194 meters, and no record of its occurrence

at the surface is known to me.

TYPE-LOCALITY.—Kagoshima Wan, Kyushu, Japan.

DISTRIBUTION.—Off the coasts of China, Korea,

and the Japanese islands of Kyushu, Shikoku, and

Honshu, except northern end of latter, to a depth

of 194 meters. The occurrence of this species in 25

meters southeast of Singapore (Boone, 1935)

seems to be considerably beyond its usual range.

REMARKS.—There is no reason to believe that

what might be construed as a dorsal lappet near

the anterior end of the sixth abdominal somite

in the illustration given by Bate (1888) is other

than an artist's error. In his description, Bate referred to this structure only as "a small tooth or.

tubercle," and the specimen came from Osaka

Wan, where L. (L.) gracilis should be expected.

Although the type-locality of L. pellucida is

well beyond the limits of the recorded range of

L. (L.) gracilis, there is little doubt that it is a

synonym of the latter species. Boone (1935) does

not compare her unique ovigerous female with

any other species, and her description and figures

do not reveal any characters that might distinguish it from L. (L.) gracilis, which is one of only

two known species in which the carapace is not

tricarinate in breeding females and the only species having a posterior spine and an otherwise entire dorsal margin on the fifth abdominal somite.
NUMBER 222 15

4. Leptochela (Leptochela) hawaiiensis,

new species

FIGURES 11-13

Leptochela robusta.—Rathbun, 1906:929. [Not L. robusta

Stimpson, I860.]

DIAGNOSIS.—Rostrum with dorsal margin variable, straight, convex, concave, or sinuous. Carapace with 3 longitudinal dorsal ridges in breeding

females only. Orbital margin minutely serrate

dorsolaterally, without mesially directed tooth on

ventral portion; suborbital angle unarmed. Fifth

abdominal somite entire, without dorsal elevations

or posterior tooth. Telson with 1 pair of dorsolateral spines in addition to anterior mesial pair;

posterior margin usually with pair (frequently

fused) of minute mesial spines in addition to

usual 5 pairs of prominent spines. Antennal scale

more than i/j> but less than 2/3 as long as carapace.

First pereopod with 21 to 30 spines on opposable

margin of movable finger. Second pereopod with

25 to 36 spines on opposable margin of movable

finger. Third pereopod with exopod not nearly

reaching distal end of ischium. Endopod of 1st

pleopod of male faintly obtuse but not flared distolaterally. Appendix masculina, not including

spines, not overreaching appendix interna. Maximum carapace length 3.0 mm.

DESCRIPTION.—Rostrum (Figure lla,&) with

dorsal margin variable, straight, convex, concave,

or sinuous, rarely overreaching eyes. Carapace

usually without median or dorsolateral carinae in

males and nonbreeding females, dorsally tricarinate over most of length in breeding females.

Orbital margin (Figure 13a,&) minutely serrate

dorsolaterally, ventral portion without mesially

directed tooth; suborbital angle rounded.

Abdomen (Figure lie) usually rounded dorsally on all somites, rarely with suggestion of blunt

carina on 5th somite. Fifth somite entire, without

dorsal prominences or posterodorsal tooth. Sixth

somite fully twice as long as high, bearing usual

transverse swelling near anterior end of dorsal

surface, slender slightly curved spine on ventrollateral surface, and distinct acute tooth on posk

FIGURE 11.—Leptochela (Leptochela) hawaiiensis, holotype, female: a, carapace and anterior

appendages; b, anterior part of carapace and rostrum, lateral aspect; c, abdomen; d, posterior

end of 6th abdominal somite; e, telson and uropods; /, posterior margin of telson; g, same,

mesial spines; h, left antennulc, dorsal aspect; i, left antenna, ventral aspect. (Magnifications:

a, c, X 12; b, e, h, i, X 25; d, X 45; /, X 62; g, X 260.)
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 12.—Leptochela (Leptochela) hawaiiensis, holotype, female: a, left and right mandibles,

dorsal (oral) aspect; b, right 1st maxilla; c, right 2nd maxilla; d, right 1st maxilliped; e, right

2nd maxilliped; /, right 3rd maxilliped; g, right 1st pereopod; h, same, fingers; i, right 2nd

pereopod; ;', same, fingers; k, right 3rd pereopod; I, right 4th pereopod; m, right 5th pereopod.

(Magnifications: /, g, i, k-m, X 25; a-e, h, j , X 62.)

terodorsal margin of posterolateral lobe (Figure

lid). Telson (Figure lie) about ly2 times as

long as 6th somite, about 2% times as long as

wide, armed with 1 pair of dorsolateral spines

slightly posterior to midlength, in addition to anterior mesial pair; posterior margin (Figures 11/,

13c) bearing pair (frequently fused, very rarely

absent) of minute mesial spines (Figures llg,

13d, i) between bases of mesial pair of usual 5

pairs of prominent spines.

Eye (Figure 13a) with papilla on mesial surface

of stalk at juncture with cornea, cornea little if

at all wider than stalk.

Antennular peduncle (Figure 11 h) with stylocerite not reaching so far as distolateral margin

of basal segment; 2nd segment about as long as

distal segment in ventromesial aspect but much

shorter in dorsal aspect.
NUMBER 222 17

FIGURE 13.—Leptochela (Leptochela) hawaiiensis, paratype,

male from Albatross Sta. 3921, carapace length 2.9 mm: a,

anterior part of carapace and eyes, dorsal aspect; b, rostrum

and orbital margin, dorsal aspect; c, posterior margin of

telson; d, same, mesial spine; e, right 1st pleopod; /, same,

endopod; g, right 2nd pleopod; h, same, appendix masculina

and appendix interna. Paratype, male from Albatross Sta.

3921, carapace length 2.8 mm: i, mesial posterior spine of

telson. (Magnifications: e, g, X 12; a, X 25; b, c, f, h, X 62;

d, i, X 260.

Antennal scale (Figure Hi) 0.5 to 0.6 as long as

carapace, 3.6 to 4.6 times as long as wide, lateral

margin concave at about midlength, blade forming rather distinct shoulder at base of distal tooth.

Distal segments of antennal peduncle about % as

wide as scale, falling considerably short of midlength of scale; ventral tooth on basal segment

neither prominent nor very sharp.

Mouthparts as illustrated (Figure 12a-/). Third

maxilliped (Figure 12/) rarely reaching as far as

distal end of antennal scale, distal segment nearly

s/4 as long as penultimate segment.

First pereopod (Figure 12g) rarely overreaching antennal scale by more than length of fingers;

fingers 1.2 to 1.5 times as long as palm; dactyl

(Figure 12ft) armed with 21 to 30 spines on opposable margin. Second pereopod (Figure 12i)

rarely overreaching antennal scale by as much

as length of fingers; fingers 1.4 to 2.0 times as long

as palm; dactyl (Figure 12/) armed with 25 to 36

spines on opposable margin. Third pereopod

(Figure 12&) overreaching extreme anterior margin of carapace by length of dactyl and about 2/5

of propodus; exopod not nearly reaching distal

end of ischium; ischium bearing row of stout setae

near extensor margin and 3 slender spines near

flexor margin; merus armed with about 5 slender

spines near flexor margin; dactyl distinctly shorter

than propodus. Fourth pereopod (Figure 12/)

reaching nearly to distal end of ischium of 2nd

pereopod when both extended anteriorly; dactyl

usually shorter than propodus. Fifth pereopod

(Figure 12m) similar to and little shorter than

4th, reaching to about midlength of ischium of

2nd pereopod when both extended anteriorly;

dactyl usually shorter than propodus.

Endopod of 1st pleopod of male (Figure l$e,f)

faintly obtuse but not flared distolaterally. Appendix masculina (Figure 13g,/») bearing 5 long

spines and not overreaching appendix interna,

without including spines. Lateral branch of upopod (Figure lie) armed with 5 to 11 movable

spines, in addition to setae.

SIZE.—Carapace lengths of males, 2.6-3.0 mm;

of nonbreeding females, 2.5-2.8 mm; of nonovigerous breeding females, 2.2, 2.4 mm; of ovigerous

females, 2.4, 2.7 mm.

MATERIAL.—HAWAII. South coast of Oahu; Diamond Head

Light, S 63°, E 1.3'; 12 m; coral and coral sand; 23.3°C (surface); 27 March 1902; 7:30 P.M.; electric light and dip net at

surface at night anchorage; Albatross Sta. 3812: 4,$ (2.6-2.8)

1 $ (2.7). Same; Diamond Head Light, S 62°, E i&; 24 m;

coral sand and broken shell; 24°C (surface); 6 May 1902;

8:45 P.M.; electrical light and dip net at surface; Albatross

Sta. 3921: 36,$ (2.6-3.0) 17$ (2.2-2.8) (2 ovig. (2.4-2.7)),

1 $ (2.2) is holotype. South coast of Molokai; Avalu Point,

Lanai, S 3/4'; 37 m; hard sand; 24°C (surface); 1 April 1902;

7:40-8:10 P.M.; electric light and dip net or surface tow net;

Albatross Sta. 3829: 2$ (2.6, 2.7).

HABITAT.—All of the specimens were taken at

the surface in early evening over bottoms of coral,
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

coral sand, broken shell, or hard sand in depths of

12 to 37 meters.

TYPE-LOCALITY.—Off Honolulu Harbor entrance, Oahu, Hawaii; surface at night over 24

meters.

DISPOSITION OF TYPES.—The female holotype

(USNM 30750) and the rest of the type-series are

deposited in the National Museum of Natural

History, Smithsonian Institution, under the catalog numbers of the United States National Museum (USNM).

DISTRIBUTION.—Known only from the type-series

collected at the surface at night south of Oahu

and south of Molokai, Hawaii, over depths of 12

to 37 meters.

ETYMOLOGY.—From Hawaii + -ensis (L., denoting locality).

REMARKS.—This species bears a close resemblance to L. (L.) bermudensis from the western

Atlantic. Males are readily distinguished from that

species by the different shape of the endopod of

the first pleopod and by the much shorter and

less spinose appendix masculina on the second

pleopod. Females differ from those of L. (L.) berFIGURE 14.—Leptochela {Leptochela) irrobusta, holotype, ovigerous female: a, carapace and

anterior appendages; b, rostrum and orbit, lateral aspect; c, same, dorsolateral aspect; d, abdomen; e, posterior end of 6th abdominal somite; /, telson and uropods; g, posterior margin of

telson; h, right antennule, dorsal aspect; i, right antenna, ventral aspect. (Magnifications:

a, d, X 12; b, c, e, f, h, i, X 25; g, X 62.
NUMBER 222 19

mudensis only in the more slender rostrum (in

dorsal aspect) and the extremely minute serrations

on the dorsolateral margin of the orbit.

Leptochela (L.) hawaiiensis superficially resembles L. (L.) irrobusta, which is common throughout the Indo-Pacific region westward from the

Marshall and Samoa islands, but the latter species

is differentiated by the extra pair of dorsolateral

spines on the telson, as well as by the mesially

directed tooth on the ventral portion of the orbital margin.

5. Leptochela (Leptochela) irrobusta, new species

FIGURES 14-18

Leptochela robusta.—Balss, 1915:17.—De Man, 1916:148

[part]; 1920:19 [part], pi. 4: fig. 7x.—Balss, 1921:7—Kemp,

1925:252.—Estampador, 1937:491.—Holthuis, 1953:52.—

Chace, 1955:5. [Not L. robusta Stimpson, I860.]

'eLeptochela robusta.—De Man, 1902:902.—Borradaile, 1917:

398.—Caiman, 1939:188.—Armstrong, 1941:1.—George and

Paulinose, 1973:196, figs. 1-5.

DIAGNOSIS.—Rostrum with dorsal margin usually straight or concave. Carapace with 3 longitudinal dorsal ridges in breeding females only. Orbital margin entire, rarely minutely serrate, with

mesially directed tooth on ventral portion; suborbital angle unarmed. Fifth abdominal somite

entire, without dorsal elevations or posterior

tooth. Telson with 2 pairs of dorsolateral spines

in addition to anterior mesial pair; posterior margin without pair of minute mesial spines in addition to usual 5 pairs of prominent spines in adults.

Antennal scale nearly 2/3 as long as carapace.

First pereopod with 12 to 40 spines on opposable

margin of movable finger. Second pereopod with

16 to 45 spines on opposable margin of movable

finger. Third pereopod with exopod not nearly

reaching distal end of ischium. Endopod of 1st

pleopod of male rounded distolaterally, not flared.

Appendix masculina, not including spines, falling

slightly short of or barely overreaching appendix

interna. Maximum carapace length 5.1 mm.

DESCRIPTION.—Rostrum (Figure I4a,b) variable,

dorsal margin usually straight or concave, rarely

slightly sinuous or convex, frequently overreaching eyes, rarely extending to level of distal margin

of basal segment of antennular peduncle. Carapace with or without obscure median dorsal carina on anterior y10 to i/3 of length in males and

nonbreeding females, tricarinate in breeding females, but lateral carinae often rounded in posterior half of length. Orbital margin (Figure 17a)

usually entire, rarely microscopically serrate,

especially in juveniles (Figure 16a); suborbital

angle rounded, but ventral margin of orbit armed

with mesially directed tooth visible only in dorsolateral view (Figure 14c).

Abdomen (Figure \4d) regularly rounded dorsally on 4 anterior somites. Fifth somite rounded

or obscurely carinate, rarely distinctly so; dorsal

margin entire, without elevations of any kind; posterior margin unarmed. Pleura of 5 anterior somites rounded in adults, those of 3rd, 4th, and

5th somites bearing small tooth in juveniles (Figure 166). Sixth somite about 1.9 times as long as

high, with usual transverse swelling and carina

near anterior end of dorsal surface, curved slender

spine posteriorly on ventrolateral surface, and distinct acute tooth on posterodorsal margin of posterolateral lobe (Figure 14e). Telson (Figure

14/), not including posterior spines, about U/2

times as long as 6th somite, about 3y3 times as

long as wide, normally armed with 2 pairs of

dorsolateral spines varying slightly in size and

position, in addition to anterior dorsomesial pair;

posterior margin (Figure 14g) without minute

pair of mesial spines between bases of mesial pair

of usual 5 pairs of prominent spines, except occasionally in juveniles (Figure 16c,d).

Eye (Figure 17a) with papilla on mesial surface of stalk proximal to cornea, cornea slightly

wider than stalk, increasing in relative diameter

with size.

Antennular peduncle (Figure I4h) with stylocerite reaching nearly as far as distolateral margin

of basal segment; 2nd segment subequal in length

to distal segment in mesial aspect, less than i/2 as

long in dorsal aspect. Dorsolateral flagellum at

least 3 times as long as carapace; ventromesial

flagellum about \y3 times as long as carapace.

Antennal scale (Figure I4i) slightly more than

0.6 as long as carapace, 3.5 to 4.6 times as long as

wide, lateral margin sinuous, distal tooth obscurely separated from mesial margin of blade.

Distal segments of antennal peduncle about 2/3 as

wide as scale, not reaching midlength of scale.

Flagellum at least 3 times as long as carapace.

Mouthparts as illustrated (Figure 15a-/). Third

maxilliped (Figure 15/) very rarely overreaching
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

m

FIGURE 15.—Leptochela (Leptochela) irrobusta, holotype, ovigerous female: a, right mandible;

b, right 1st maxilla; c, right 2nd maxilla; d, right 1st maxilliped; e, right 2nd maxilliped;

/, right 3rd maxilliped; g, right 1st pereopod; h, same, fingers; i, right 2nd pereopod; /', same,

fingers; k, right 3rd pereopod; /, right 4th pereopod; m, right 5th pereopod. (Magnifications:

c-g, i, k-m, X 25; a, b, h, j , X 62.)
NUMBER 222 21

FIGURE 16.—Leptochela (Leptochela) irrobusta, paratype,

juvenile from Bikini lagoon, carapace length 1.7 mm: a,

rostrum and dorsal orbital margin; b, abdomen; c, posterior

margin of telson; d, same, mesial spines. (Magnifications:

b, X 25; a, c, X 62; d, X 260.)

antennal scale, distal segment %0 to % as long

as penultimate segment.

First pereopod (Figure 15g) overreaching antennal scale, occasionally by as much as length of

fingers; fingers 1.3 to 2.0 times as long as palm;

dactyl (Figure \5h) armed with 12 to 40 spines

on opposable margin. Second pereopod (Figure

15i) rarely overreaching antennal scale by more

than length of fingers; fingers 1.5 to 2.2 times as

long as palm; dactyl (Figure 15;) armed with 16

to 45 spines on opposable margin. Third pereopod (Figure 15k) overreaching extreme anterior

margin of carapace by about length of dactyl; exopod falling far short of distal end of ischium;

ischium armed with row of about 4 stout spines

near flexor margin; merus with about 5 longer

stout spines near flexor margin; dactyl variable in

length and shape, usually shorter than propodus.

Fourth pereopod (Figure 15/) reaching to about

distal y5 of ischium of second pereopod when

both extended anteriorly; dactyl usually longer,

occasionally slightly shorter than propodus. Fifth

pereopod (Figure 15m) similar to and nearly as

long as 4th, reaching at least midlength of ischium of 2nd pereopod when both extended anteFIGURE 17.—Leptochela (Leptochela) irrobusta, paratype,

male from Bikini lagoon, carapace length 3.6 mm: a, anterior

part of carapace and eyes, dorsal aspect; b, right 1st pleopod; c, same, endopod; d, right 2nd pleopod; e, same,

appendix masculina and appendix interna. (Magnifications:

b, d, X 12; a, X 25; c, e, X 62.)

riorly; dactyl usually slightly shorter than propodus.

Endopod of first pleopod of male (Figure 176,

c) subelliptical, fully 3 times as long as wide. Appendix masculina (Figures \7d,e, \%a,b) bearing

3 to 7 long spines, usually in distal half, falling

slightly short of or extending very little beyond

end of appendix interna, not including spines.

Lateral branch of uropod (Figure 14/) armed

with 8 to 16 movable spines, in addition to setae.

SIZE.—Carapace lengths of males, 1.8-5.1 mm;

of nonbreeding females, 2.0-5.1 mm; of nonovigerous breeding females, 3.2-4.3 mm; of ovigerous

females, 2.7-4.6 mm. The smallest male has a rudimentary appendix masculina, but appendices

masculinae may be less than fully developed in

some males with a carapace length up to 3.0 mm.

MATERIAL.—PERSIAN GULF. About 200 m off southwest

end of Juraid Island; 27°10'N, 49C51'E; 9 m; massive coral

formations interspersed with patches of coral sand; 5 October 1956; 7:45 A.M.; Petersen sample; C. E. Dawson Sta. 7:

13 (2.9).
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 18.—Leptochela (Leptochela) irrobusta: a, appendices

masculinac of male paratypes selected at random from

Bikini lagoon, with carapace lengths (mm) as indicated;

b, appendix masculina of male paratype, carapace length

5.0 mm, from Albatross Sta. 5134. (Magnifications: a, b,

X 62.)

VIETNAM. Bay of Nha Trang; 1 January to 10 April

1960; Ariel Gallardo Sta. 220a: \$ (4.3). Same; Sta. 317:

1$ (3.6).

PHILIPPINE ISLANDS. Sablayan Anchorage, Mindoro;

electric light; 13 December 1908; Albatross: 1 $ (3.1). Batan

Island, Lagonoy Gulf; electric light; 5 June 1909; Albatross:

1 ovig. 9 (4.0). Port Cataingan, Masbate; electric light; 19

April 1908; Albatross: 1 $ (2.8). Nogas Point, Panay; electric

light; 3 February 1908; Albatross: 10 $ (3.1-3.7) 9$ (2.8-

3.8) 4 juv. (1.7-2.3). Mahinog Anchorage, Camiguin Island;

electric light; 3 August 1909; Albatross: 2$ (2.5, 2.6) 9$

(3.0-3.9). Sulu Sea off Panabutan Point, Mindanao; 49 m;

green mud, coral sand; 6 February 1908; 9:27 A.M.; 9-foot

Tanner beam trawl, mud bag; Albatross Sta. 5131: 1 $ (5.0).

Same; 70 m; green mud, sand; 6 February 1908; 10:40 A.M.;

9-foot Tanner beam trawl, mud bag; Albatross Sta. 5133:

4$ (4.7-5.1) 10$ (4.2-5.1) (7 ovig. (4.2-4.6)). Off Balukbaluk Island, near fiasilan Island, Sulu Archipelago; 6°44'45"

N, 121 °48' E; 46 m; fine sand; 7 February 1908; 7:22 A.M.;

9-foot Tanner beam trawl, mud bag; Albatross Sta. 5134: 1 $

(5.0). Off Jolo Island, Sulu Archipelago; 6°06'N, 120°58'50"

E; 35 m; sand, coral; 14 February 1908; 10:55 A.M.; 12-foot

Agassiz beam trawl two mud bags; Albatross Sta. 5138: 1

ovig. $ (4.4).

INDONESIA. Labuan Blanda Island, Selat Butung, Celebes; electric light; 13 December 1909; Albatross: 1$ (3.6)

2$ (3.6, 3.7).

MARSHALL ISLANDS. Eniwctok Atoll; lagoon; subtidal;

6 June 1946; M. W. Johnson: 1 $ (3.5). Eniwetok Atoll;

lagoon off Eniwetok Island at Bowditch anchorage; surface

light at night; 23 May 1946; L. P. Schultz: 2$ (2.3) 3$

(2.2-2.3) 4 juv. (1.8-2.0). Eniwetok Atoll; Southwest Passage,

leeward side of reef 3.7 km south of Rigili Island; surface

light at night; 24 May 1946; L. P. Schultz: 10 $ (2.1-3.1)

18$ (2.0-3.3) 10 juv. (1.6-1.9). Bikini Atoll; northeast end

of lagoon at Bowditch anchorage; surface light at night; 30

March 1946; L. P. Schultz and V. E. Brock: 42 $ (2.0-3.2)

83$ (2.0-3.4) (8 ovig. (2.7-3.2)) 16 juv. (1.7-2.0). Same; 23

April 1946; L. P. Schultz and J. P. E. Morrison: 46,$ (2.0-

3.1) 46$ (2.1-3.7) 8 juv. (1.8-2.0) + 4524, $, & juv. (1

ovig.). Same; 24 April 1946; L. P. Schultz: 47 $ (1.8-3.4) 46$

(2.2-3.4) (5 ovig. (2.9-3.2)) 7 juv. (1.7-2.1) + 1,731 ^, $,

& juv. (13 ovig.), 1 ovig. $ (3.0) is holotype. Same; 25

April 1946; L. P. Schultz: 40 $ (2.0-3.4) 51 $ (2.0-3.5) 9 juv.

(1.4-1.9) + 1,005 <$, $, & juv. (4 ovig.). Bikini Atoll; lagoon, 7.4 km south of west end of Bikini Island; about 45 m;

sand and Halimeda; 26 August 1947; J. P. E. Morrison: 1 $

(2.8). Rongelap Atoll; lagoon, 0.9 km off Yugui Island; 24 m;

surface light at night; 30 July 1946; E. S. Herald: 20^ (2.0-

3.0) 36$ (2.0-3.0) 74 juv. (1.4-2.0). Rongelap Atoll; lagoon,

0.9 km off Lomuilal Island; 24 m; surface light at night; 31

July 1946; E. S. Herald: 2<J (2.3) 3$ (2.1-2.9). Rongerik

Atoll; lagoon, 183 m off F.niwetak Island; surface light at

night; 28 June 1946; L. V. Schultz and E. S. Herald: 12 $

(2.4-3.4) 5$ (2.5-3.1). Ujae Atoll; 60 m outside Ujae Passage; surface at stern of ship under lights; 1951-1952; F. S.

MacNeil: 5^ (2.7-3.0) 3$ (2.0-3.1).

HABITAT.—It will be noted that 2/3 of the lots

and all but 24 of the 3876 specimens listed above

were taken at surface lights at night. That the species sometimes occurs on or near the bottom, however, is indicated by the 14 specimens trawled at

Albatross station 5133 and, especially, by the single male taken in a Petersen grab in the Persian

Gulf. The trawled specimens were found in depths

of 35 to 70 meters on bottoms of mud or sand,

sometimes with fragments of coral or calcareous

algae.

TYPE-LOCALITY.—Lagoon of Bikini Atoll, Marshall Islands; surface at night.

DISPOSITION OF TYPES.—The ovigerous female

holotype (USNM 94729) and the rest of the typeseries are deposited in the National Museum of

Natural History, Smithsonian Institution, under

the catalog numbers of the United States National

Museum (USNM).

DISTRIBUTION.—Red Sea and Persian Gulf, Laccadive, Maldive, Andaman, and Nicobar islands,

Burma, Western Australia, Indonesia, and Philippine Islands eastward to the Marshall and Samoa

islands, at the surface and to depths of 118 and
NUMBER 222 23

possibly 275 meters. (See L. (L.) robusta.)

ETYMOLOGY.—From ir- (L., = not) + robusta.

REMARKS.—Quite understandably, this abundant

wide-ranging, and variable little species has been

confused heretofore with L. (L.) robusta. Immature females of the latter species can seldom be

distinguished from L. (L.) irrobusta with confidence. In general, L. (L.) irrobusta has the rostrum proportionately longer, the spine on the ventrolateral surface of the posterior part of the

sixth abdominal somite curved rather than nearly

straight, and the posterior pair of dorsolateral

spines arising near the midlength of the telson

rather than more posteriorly, but the species is so

variable that these characters are not always reliable. The size of sexually mature individuals—

carapace lengths of 2.7 to 4.6 mm in L. (L.) irrobusta and more than 6 mm in L. (L.) robusta—

is important, but it is useful only for tricarinate

females, because nonbreeding females may be

larger than ovigerous ones. The only really definitive character lies in the length, form, and spination of the male appendix masculina (compare

Figures 18 and 2Se-h). That this character is

meaningful is confirmed by comparing Figure 28/

showing the form of the appendix masculina in

a male L. (L.) robusta with a carapace length of

3.0 mm, with Figure 18&, which illustrates the appendix in a considerably larger male of L. (L.)

irrobusta.

Most of the Red Sea specimens examined

through the kind cooperation of L. B. Holthuis

and mentioned in the Remarks section under L.

aculeocaudata belong to L. irrobusta, but they

are not otherwise treated as part of this report nor

are they included in the type-series of this species.

6. Leptochela (Leptochela) japonica

Hayashi and Miyake

FICURES 19-21

Leptochela japonica Hayashi and Miyake, 1969:1, figs. 1, 2.—

Fujino and Miyake, 1970:242.

DIAGNOSIS.—Rostrum with dorsal margin slightly

sinuous. Carapace with 3 longitudinal dorsal

ridges in both males and females. Orbital margin

entire, not serrate, without mesially directed tooth

on ventral portion; suborbital angle unarmed.

Fifth abdominal somite with 3 distinct elevations

in dorsal midline, posterior margin bluntly produced mesially. Telson with 1 pair of dorsolateral

spines in addition to anterior mesial pair; posterior margin without pair of minute mesial spines

in addition to usual 5 pairs of prominent spines.

Antennal scale slightly more than i/2 as long as

carapace. First pereopod with about 48 spines on

opposable margin of movable finger. Second pereopod with about 60 spines on opposable margin of

movable finger. Third pereopod with exopod not

nearly reaching distal end of ischium. Endopod

of 1st pleopod of male not flared distolaterally.

Appendix masculina, not including spines, slightly

overreaching appendix interna. Maximum carapace length 6.2 mm.

DESCRIPTION OF MALE.—Rostrum (Figure 19a)

with dorsal margin faintly sinuous, not overreaching eyes. Carapace with distinct median carina

extending posteriorly slightly beyond midlength

and with well-marked pair of dorsolateral carinae

extending to about posterior 1.3 of carapace. Orbital margin entire, not serrate, ventral portion

without mesially directed tooth; suborbital angle

broadly rounded, unarmed.

Abdomen (Figure 19&) rounded dorsally on 3

anterior somites, distinctly carinate in midline on

4th and 5th, latter surmounted by 3 low but distinct elevations and projecting posteriorly as blunt

tooth. Sixth somite about I14 times as long as

high, transverse swelling near anterior end of dorsal surface bearing high sharp carina trending

slightly anteriorly; spine on ventrolateral surface

slightly curved; posterolateral lobe (Figure 19c)

without tooth on posterodorsal margin. Telson

(Figure 19d) less than \i/2 times as long as 6th

somite, about 2y2 times as long as wide, armed

with 1 pair of dorsolateral spines arising at about

midlength, in addition to anterior mesial pair;

posterior margin without pair of minute spines

between bases of mesial pair of usual 5 pairs of

prominent spines.

Eye with papilla on mesial surface of stalk proximal to juncture with cornea, cornea unusually

inflated, much wider and longer than stalk.

Antennular peduncle (Figure \9e) with stylocerite reaching about as far as distolateral margin

of basal segment; 2nd segment about as long as

distal segment in ventromesial aspect, much

shorter in dorsal aspect.

Antennal scale (Figure 19/) nearly 0.6 as long
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Leptochela (Leptochela) japonica, male from Gulf of Thailand, carapace length

5.8 mm: a, carapace and anterior appendages; b, abdomen; c, posterior end of 6th abdominal

somite; d, telson and uropods; e, right antennule, dorsal aspect; /, right antenna, ventral aspect.

(Magnifications: a, b, X 6; d-f, X 12; c, X 25.)

as carapace, 3.4 times as long as wide, lateral margin convex distally, nearly straight on proximal

2/s of length, blade forming slight shoulder at base

of distal tooth. Distal segments of antennal peduncle about %0 as wide as scale, reaching nearly

to midlength of scale; ventral tooth on basal segment blunt, rather indistinct.

Mouthparts as illustrated (Figure 20a-/). Mandible (Figure 20a) with rather distinct angle in

distal margin of incisor process. Third maxilliped

(Figure 20/) not reaching as far as distal end of

antennal scale, distal segment nearly % as long as

penultimate segment.

First pereopod (Figure 20g) overreaching antennal scale by little more than length of fingers;

fingers about as long as palm; dactyl (Figure 20/J)

armed with 48 spines on opposable margin. Second pereopod (Figure 20i) overreaching antennal

scale by nearly length of fingers; fingers 1.3 times

as long as palm; dactyl (Figure 20;) armed with

60 spines on opposable margin. Third pereopod

(Figure 20A) overreaching extreme anterior margin of carapace by nearly i/2 length of dactyl;

exopod not nearly reaching distal end of ischium;

ischium with row of about 6 slender spines near

middle of lateral surface and 3 oblique rows of 3,

8, and 7 stouter spines near flexor margin; merus

with row of 10 stout spines, increasing in length

distally, near flexor margin; dactyl subequal in

length of propodus. Fourth pereopod (Figure 20/)

reaching at least to midlength of 2nd pereopod

when both extended anteriorly; merus with rather

discrete subrectangular lobe bearing 2 stout spines

on flexor margin; dactyl shorter than propodus.
NUMBER 222 25

FIGURE 20.—Leptochela {Leptochela) japonica, male from Gulf of Thailand, carapace length

5.8 mm: a, right mandible; b, right 1st maxilla; c, right 2nd maxilla; d, right 1st maxilliped;

e, right 2nd maxilliped; /, right 3rd maxilliped; g, right 1st pereopod; h, same fingers; i, left

2nd pereopod; /', same, fingers; k, right 3rd pereopod;' /, right 4th pereopod; tn, right 5th

pereopod. (Magnifications: /, g, i, k-tn, X 12; a-e, h, j , X 25.)
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Fifth pereopod (Figure 20m) reaching to about

distal i/3 of ischium of 3rd pereopod when both

extended anteriorly; similar to 4th, especially in

form of merus, but distinctly shorter; dactyl much

shorter than propodus.

Endopod of first pleopod (Figure 21a,b) long

and comparatively narrow, nearly 3 times as long

as wide, broadly and obscurely obtuse, but not

flared, distolaterally. Appendix masculina (Figure 2lc,d) bearing 7 long spines and slightly

overreaching appendix interna, not including

spines. Lateral branch of uropod (Figure 19rf)

armed with about 20 movable spines increasing

in length distally.

SIZE.—Carapace lengths of males, 4.0 and 5.8

mm. Hayashi and Miyake (1969:5) noted that the

four ovigerous females in the type-series have carapace lengths of 5.5-6.2 mm, and the single female

without eggs has a carapace length of 4.2 mm.

MATERIAL.—GULF OF THAILAND. Shore reef region on

east side of Goh Kram Island; 12°41'35"N, 100°48'28"E; 25

October 1957; George Vanderbilt Foundation Sta. 8: 1 $ (5.8).

QUEENSLAND, AUSTRALIA. Bustard Bay (24°05'S,

151°48'E); 19 September 1938; Haul 46/38 (old number,

Net 200; K. Sheard: l £ freshly molted (4.0) (Australian

Museum).

HABITAT.—The four ovigerous females in the

type-series were dredged in 45 and 65-66 meters;

d

FIGURE 21.—Leptochela (Leptochela) japonica, male from

Gulf of Thailand, carapace length 5.8 mm: a, right 1st

pleopod; b, same, endopod; c, right 2nd pleopod; d, same,

appendix masculina and appendix interna. (Magnifications:

a, c, X 12; b, X 25; d, X 62)

the single female without eggs was taken in a surface plankton net at night.

TYPE-LOCALITY.—Tachibana-wan (Chijiwa Bay

northwestern Kyushu, Japan; 65-66 meters.

DISTRIBUTION.—Korea Strait; northeast of Twiwan; Gulf of Thailand; and Queensland, Australia; 45-111 meters.

REMARKS.—Proof of the identity of the male

from the Gulf of Thailand with L. (L.) japonica

must be deferred until additional material becomes

available. The Thailand specimen has the dorsal

projections on the fifth abdominal somite less

prominent, and there seem to be subtle and unexpected differences in some of the mouthparts,

but the specimen agrees in most other particulars

with the description of L. (L.) japonica, even to

the spination on the third pereopod and the characteristic shape of the merus of the fourth and

fifth pereopods. Certainly the most interesting

feature of the specimen is the dorsally tricarinate

carapace, a development that was previously believed to be restricted to breeding females in

Leptochela.

In the freshly molted male from Queensland,

the projections of the fifth abdominal somite are

stronger, much as depicted by Hayashi and Miyake (1969: fig. la), and the carapace—as well as

can be determined from the soft and misshapen

integument—seems to be tricarinate to the same

extent as in the male from Thailand.

There is little doubt that L. (L.) japonica is

related to L. (P.) carinata from the western Atlantic. Both species have dorsal projections on

the fifth abdominal somite, unusually numerous

spines on the opposable margins of the fingers of

the two anterior pairs of pereopods, and reduced

fifth pereopods. Leptochela (L.) japonica differs

rather strikingly from L. (P.) carinata, however,

in lacking the hinged dorsal lappet on the sixth

abdominal somite, in the form of the telson, and

in the much shorter and broader antennal scale.

The former species therefore seems to represent

a link between the two subgenera proposed above.

7. Leptochela (Leptochela) papulata, new species

FIGURES 22-24

DIAGNOSIS.—Rostrum with dorsal margin convex or slightly sinuous. Carapace with 3 longitudi-
NUMBER 222 27

FIGURE 22.—Leptochela (Leptochela) papulata, holotype, ovigerous female: a, carapace and

anterior appendages; b, anterior part of carapace and eyes, dorsal aspect; c, abdomen; d, posterior end of 6th abdominal somite; e, right antennule, dorsal aspect; /, right antenna, ventral

aspect. Paratype, ovigerous female from Grampus Sta. 5099, carapace length 4.4 mm: g, 5th

abdominal somite, lateral aspect. (Magnifications: a, c, g, X 12; b, d-f, X 25.)

nal dorsal ridges in breeding females only. Orbital

margin entire, rarely minutely serrate, without

mesially directed tooth on ventral portion; suborbital angle unarmed. Fifth abdominal somite

with 1 to 3 more or less distinct pimplelike elevations in posterior half of dorsal midline, without

posterior tooth. Telson with 1 pair of dorsolateral

spines in addition to anterior mesial pair; posterior margin with pair of minute mesial spines in

addition to usual 5 pairs of prominent spines. Antennal scale barely y2 as long as carapace. First

pereopod with 16 to 28 spines on opposable margin of movable finger. Second preopod with 22 to

36 spines on opposable margin of movable finger.

Third pereopod with exopod not nearly reaching

distal end of ischium. Endopod of 1st pleopod of

male flared distolaterally. Appendix masculina,

not including spines, distinctly overreaching appendix interna. Maximum carapace length 4.4

mm.

DESCRIPTION.—Rostrum (Figure 22a) usually

with dorsal margin regularly convex, rarely

slightly concave in anterior half, usually falling

short of distal surface of eye, rarely overreaching

basal segment of antennular peduncle. Carapace

with median dorsal carina on anterior % to i/2 of

length in males and nonbreeding females, sharply

tricarinate over most of length in breeding females. Orbital margin (Figure 226) entire, rarely

minutely serrate, ventral portion without mesially
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 23.—Leptochela (Leptochela) papulata, paratype, carinate female from Atlantis Sta.

A-266-36, carapace length 2.7 mm: a, telson and left uropod; b, posterior margin of telson;

c, same, mesial spines; d, right mandible; e, right 1st maxilla; /, right 2nd maxilla; g, right 1st

maxilliped; h, right 2nd maxilliped; i, right 3rd maxilliped; /, right 1st pereopod; k, same,

fingers; /, right 2nd pereopod; m, same, fingers; n, right 3rd pereopod; o, right 4th pereopod;

p, right 5th pereopod. (Magnifications: a, f-j, I, n-p, X 25; b, d, e, k, m, X 62; c, X 260.)
NUMBER 222 29

directed tooth; suborbital angle rounded, not

dentate.

Abdomen (Figure 22c) regularly rounded dorsally on 4 anterior somites. Fifth somite (Figure

22c,g) bluntly carinate dorsally, carina usually

surmounted in posterior half by 1 to 3 (usually

3) pimplelike elevations, rarely with barely visible

vestiges thereof; posterior margin unarmed. Sixth

somite about 1.6 times as long as high, with usual

transverse swelling and carina near anterior end

of dorsal surface, long slender spine on ventrolateral surface, and usually with minute blunt

tooth (Figure 22d) on posterodorsal margin

of posterolateral lobe (tooth sometime lacking,

rarely prominent and acute). Telson (Figure

23a), not including posterior spines, about 1.8

times as long as 6th somite, about 3.4 times as long

as wide, armed with 1 pair of dorsolateral spines

at about midlength, in addition to anterior mesial

pair; posterior margin (Figure 236) bearing pair

of minute spines (Figure 23c) between bases of

mesial pair of usual 5 pairs of prominent spines.

Eye (Figure 226) with papilla on mesial surface of stalk proximal to cornea, cornea slightly

wider than stalk.

Antennular peduncle (Figure 22e) with stylocertie reaching nearly as far as distolateral margin

of basal segment; 2nd segment longer than distal

segment in mesial aspect but distinctly shorter in

dorsal aspect.

Antennal scale (Figure 22/) barely 0.5 as long

as carapace, 2.8 to 3.5 times as long as wide, lateral

margin sinuous, blade forming rather distinct

shoulder at base of distal tooth. Distal segments

of antennal peduncle about 2/3 as wide as scale

and nearly reaching midlength of scale.

Mouthparts as illustrated (Figure 23d-t). Third

maxilliped (Figure 23i) rarely overreaching antennal scale, distal segment about % as long as

penultimate segment.

First pereopod (Figure 23;) usually overreaching antennal scale by little more than length of

fingers; fingers 1.2 to 1.5 times as long as palm;

dactyl (Figure 23*) armed with 16 to 28 spines

on opposable margin. Second pereopod (Figure

23/) sometimes overreaching antennal scale by

more than length of fingers; fingers 1.6 to 2.0 times

as long as palm; dactyl (Figure 23m) armed with

22 to 36 spines on opposable margin. Third pereopod (Figure 23n) overreaching extreme anterior

FIGURE 24.—Leptochela (Leptochela) populate, paratype,

male from Beveridge Sta. BST. 271, V, carapace length

2.6 mm: a, right 1st pleopod; b, same, endopod; c, right

2nd pleopod; d, same, appendix masculina and appendix

interna. (Magnifications: a, c, X 12; b, d, X 62.)

margin of carapace by about length of dactyl;

exopod falling far short of distal end of ischium;

ischium armed with row of about 9 slender sharp

spines near extensor margin and 3 subequally

spaced stout spines on lateral surface; merus with

4 longer blunt spines on lateral surface; dactyl

slightly shorter than propodus. Fourth pereopod

(Figure 23o) overreaching ischium of 2nd pereopod by about length of dactyl when both extended

anteriorly; dactyl shorter than propodus. Fifth

pereopod (Figure 23/)) similar to and only slightly

shorter than 4th, reaching to about distal ]/s of

ischium of 2nd pereopod when both extended anteriorly; dactyl nearly as long as propodus.

Endopod of 1st pleopod of male (Figure 24a,6)

with lateral margin flared distally. Appendix masculina (Figure 24c,d) bearing 7 long spines, distinctly overreaching appendix interna, without

including spines. Lateral branch of uropod (Figure 23a) armed with 10 to 17 movable spines, in

addition to setae.

SIZE.—Carapace lengths of males, 2.1-3.9 mm;

of nonbreeding females, 2.2-3.6 mm; of nonovigerous breeding females, 2.7, 3.9 mm; of ovigerous

females, 3.2-4.4 mm; of juveniles, 1.6-2.0 mm. The

smallest male has a barely discernible appendix

masculina; the smallest male with a fully developed appendix masculina has a carapace length

of 2.6 mm.
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MATERIAL.—NORTH CAROLINA. East of Cape Lookout;

34°35'3O"N, 75°45'30"W; 59 m; white sand with black specks;

18 October 1885; large beam trawl; Albatross Sta. 2605: 1

ovig. $ (3.8) holotype. Southeast of Cape Lookout; 34°34.5'N,

76°255'W; 20 m; sand and broken shell; 9°C; 19 January

1966; dredge; Beveridge (J. H. Day Sta. BST. 271. V.): 1 &

(2.6) 1 $ (2.5). Same locality and date; 0.2m2 Van Veen

grab; Beveridge (J. H. Day Sta. BST 274. E.): 1$ (2.2).

34°28.5'N, 76°13.1'W; 33 m; 15 February 1965; Van Veen

grab; Eastward Sta. 489: 1 juv. (2.0). 34°28.1'N, 76°20.0'W;

27 m; 15 February 1965; small biological trawl; Eastward

Sta. 497: 1 $ (2.7) 1 juv. (2.0). 34°24.8'N, 75°59.5'W; 40 m;

coarse sand and mud; 20.6°C; 30 November 1965; dredge;

Eastward Sta. 3464 (J. H. Day Sta. BST. 236. P.): ly£ (2.1).

34°20.0'N, 75°54.5'W; 202 m; hard packed; 8 January 1966;

dredge; Eastward Sta. 3538 (J. H. Day Sta. BST. 289, M.):

1 juv. (1.6 + ). South of Cape Lookout; 34°04.5'N, 75°25.5'W;

20 m; coarse sand; 20°C; 24 November 1965; 0.2m2 Van Veen

grab; Beveridge (J. H. Day Sta. BST. 233. N.): 1 ovig. $

(3.5).

SOUTH CAROLINA. Southeast of Charleston; 32°12'N,

79°15'W; 46 m; terigenous sand, shell debris; 12 July 1961

(1730-1732); Van Veen grab; Atlantis Sta. A-266-36: 1 ?

(2.7).

GEORGIA. Sapelo Whistle 335°-5i/2 km; 21 m; 28 February 1962; M. Gray: 1 $ (3.6). East of Doboy Sound Sea Buoy

13-46 km; 16-22 m; 4 December 1962; M. Gray: 1 $ (2.9).

GULF OF MEXICO. South of Cape San Bias, Florida;

29°15230"N, 85°29'30"W; 49 m; gravel; 7 February 1885;

large beam trawl; Albatross Sta. 2372: 1 $ (3.6). Anclote

Light, Florida, E 7/8 N-39.8 km; 23 m; rock, coral, and sand;

17.2°C; 28 March 1901 (10:30 A.M.); dredge; Fish Hawk Sta.

7106: 1 $ (32). Southwest of Sarasota, Florida; 27°04'00"N,

83°21'15"W; 48 m; coarse gray sand and broken shell; 18

March 1885; large beam trawl; Albatross Sta. 2409: 1 $ (3.9).

West of Captiva Island, Florida; 26°33'N, 83°1(KW; 51 m;

sandy; 19.1 °C; 2 April 1901 (6.30 A.M.); 7-ft beam trawl;

Fish Hawk Sta. 7123: 1 ovig. $ (3.9). West of Marco Florida;

26°04'00"N, 82°49'00"W; 39 m; sand and broken shell; 17

March 1889; hand dredge; Grampus Sta. 5099: 1 ovig. $

(4.4). 25°54'OO"N, 83°20'00"W; 57 m; gray sand with black

specks and broken shell; 15 March 1889; hand dredge;

Grampus Sta. 5092; 2 9 (3.2, 3.9) (1 ovig. (3.2)). Northwest

of Dry Tortugas; 25°34'OO"N, 83°28'0O"W; 71 m; gray coral

and fine shell; 1 March 1889; hand dredge; Grampus Sta.

5076: 13 (3.8).

HABITAT.—In depths of 20 to 202 meters, usually

less than 70 meters, commonly on bottoms of sand,

frequently with shell fragments, occasionally on

gravel, rock, and coral, at bottom temperatures

ranging from 9° to 20.6°C.

There is no evidence from the material available that L. (L.) papulata approaches the surface

at night, as do L. (L.) bermudensis and L. (L.)

serratorbita, but additional surface or subsurface

collections in the region inhabited by the former

species may refute this assumption; the fact that,

on four different occasions, specimens of L. (L.)

papulata were found in grab samples indicates

that the species must spend some time in actual

contact with the bottom.

TYPE-LOCALITY.—East of Cape Lookout, North

Carolina; 34°35'30"N, 75°45'30"W; 59 meters.

DISPOSITION OF TYPES.—The ovigerous female

holotype (USNM 23375) and the rest of the typeseries are deposited in the National Museum of

Natural History, Smithsonian Institution, under

the catalog numbers of the United States National

Museum (USNM).

DISTRIBUTION.—Off North Carolina to Georgia

and the eastern Gulf of Mexico, in depths of 20

to 202 meters.

ETYMOLOGY.—From papula (L., = pimple) +

atus (L., = provided with), in reference to the

one to three elevations usually present in the dorsal midline of the fifth abdominal somite.

REMARKS.—Apparently this species is closely related to L. (L.) bermudensis, as indicated by the

form of the endopod of the first pleopod in males

and the length of the appendix masculina, as well

as by most other characters. It may be a slightly

larger species, the smallest ovigerous female having a carapace length of 3.2 mm, whereas the largest specimens of L. (L.) bermudensis currently

available have a carapace length of only 3.0 mm,

and females of that species may be ovigerous at a

carapace length of 2.4 mm. Most specimens of

L. papulata may be distinguished by the one to

three (usually three) elevations in the dorsal

midline of the fifth abdominal somite; occasionally, even in mature individuals, these elevations

are so obscure as to be barely discernible, but the

dorsal margin of the fifth somite is always faintly

sinuous in lateral view, rather than regularly convex as in L. (L.) bermudensis. Also, the posterolateral extension of the sixth abdominal somite

is usually unarmed or bears an obscure blunt or

subacute tooth, rather than the prominent acute

tooth usually present in L. (L.) bermudensis;

however, occasional specimens of L. (L.) papulata

have a tooth nearly as prominent and sharp as

in L. (L.) bermudensis. Finally, the anterior margin of the carapace is usually convex in L. (L.)

papulata, whereas it is often concave in L. (L.)

bermudensis. A few of the specimens of /,. (L)

papulata examined have the dorsolateral margin
NUMBER 222 31

of the orbit minutely serrate, but these serrations

are even more obscure than in L. (L.) haioaiiensis and much less prominent than in L. (L.) serratorbita.

8. Leptochela (Leptochela) pugnax De Man

FICURES 25-27

Leptochela pugnax De Man, 1916:148; 1920:26, pi. 4: fig. 8.—

Kemp, 1925:255.—Armstrong, 1941:3— Kubo, 1955:101, figs.

4, 5.—Holthuis and Gottlieb, 1958:22.

^Leptochela sp. De Man, 1920:30.

Leptochela robusta.—Miyzdi, l940b:141; 1941:180. [Not L.

robusta Stimpson, I860.]

DIAGNOSIS.—Rostrum with dorsal margin concave. Carapace with 3 longitudinal dorsal ridges

in breeding females only. Orbital margin entire,

not serrate, without mesially directed tooth on

ventral portion; suborbital angle distinctly dentate.

Fifth abdominal somite entire, without dorsal elevations or posterior tooth. Telson with 1 pair of

dorsolateral spines in addition to anterior mesial

pair; posterior margin without pair of minute

mesial spines in addition to usual 5 pairs of prominent spines. Antennal scale more than 1/2 but

usually less than 2/3 as long as carapace. First pereopod with 16 to 21 spines on opposable margin

of movable finger. Second pereopod with 17 to 26

spines on opposable margin of movable finger.

Third pereopod with exopod not nearly reaching

distal end of ischium. Endopod of first pleopod of

male slightly but not conspicuously flared distolaterally. Appendix masculina, not including

spines, not overreaching appendix interna. Maximum carapace length 3.4 mm.

DESCRIPTION.—Rostrum (Figure 25a,b) distinctly

upturned, dorsal margin concave, rarely reaching

as far as distal end of basal segment of antennular peduncle. Carapace with sometimes obscure

median dorsal carina on anterior 14 to i/3 of

length but without paired dorsolateral ridges in

males and nonbreeding females, median carina

on y4 to %0 of length and paired dorsolateral

ridges on ]/2 to % of length in breeding females.

Orbital margin (Figure 256) entire, not serrate,

ventral portion without mesially directed tooth;

suborbital angle distinctly dentate.

Abdomen (Figure 25c) regularly rounded dorsally on 3 anterior somites, occasionally bluntly

FIGURE 25.—Leptochela (Leptochela) pugnax, ovigerous female from Nicobar Islands, carapace length 2.5 mm: a, carapace and anterior appendages; b, rostrum and orbit, lateral

aspect; c, abdomen; d, posterior end of 6th abdominal

somite; e, telson and uropods; /, posterior margin of telson;

g, same, mesial part. (Magnifications: a, c, X 12; b, d, e,

X 25; /, X 62; g, X 260.)

and inconspicuously carinate on 4th and 5th.

Fifth somite without dorsal prominences and

without posterodorsal tooth. Sixth somite about

1.8 times as long as high, anterodorsal transverse

swelling not prominent, spine on ventrolateral surface curved and distinct but not very long, posterolateral lobe bearing distinct acute tooth on posterodorsal margin (Figure 25d). Telson (Figure

25e), not including posterior spines, about 1.3

times as long as 6th somite, slightly less than 3
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 26.—Leptochela (Leptochela) pugnax, male from Nicobar Islands, carapace length

2.8 mm: a, right antennule, dorsal aspect; b, right antenna, dorsal aspect; c, right mandible;

d, right 1st maxilla; e, right 2nd maxilla; /, right 1st maxilliped; g, right 2nd maxilliped;

h, right 3rd maxilliped; i, right 1st pereopod; ;*, same, fingers; k, right 2nd pereopod; I, same,

fingers; m, right 3rd pereopod; n, right 4th pereopod; o, right 5th pereopod. (Magnifications:

a, b, h, i, k, m-o, X 25; c-g, j , I, X 62.)
NUMBER 222 33

times as long as wide, armed with 1 pair of dorsolateral spines slightly anterior to midlength, in

addition to anterior mesial pair; posterior margin

(Figure 25f,g) without pair of minute spines between bases of mesial pair of usual 5 pairs of

prominent spines.

Eye with rather prominent papilla on mesial

surface of stalk proximal to cornea, cornea little

wider than stalk.

Antennular peduncle (Figure 26a) with stylocerite reaching nearly as far as distolateral margin

of basal segment; 2nd segment about as long as

distal segment in mesial aspect, distinctly shorter

in dorsal aspect.

Antennal scale (Figure 266) 3.7 to 4.6 times as

long as wide, lateral margin concave, blade only

occasionally forming suggestion of subdistal shoulder, distal tooth usually continuous with mesial

margin of blade. Distal segments of antennal peduncle nearly as wide as scale, reaching nearly to

midlength of scale; basal segment without ventral

tooth.

Mouthparts as illustrated (Figure 26c-h). Third

maxilliped (Figure 26h) not reaching as far as

distal end of antennal scale, distal segment at least

% as long as penultimate segment.

First pereopod (Figure 26£) rarely overreaching antennal scale by as much as length of fingers;

fingers 1.5 to 2.0 times as long as palm; dactyl

(Figure 26;) armed with 16 to 21 spines on opposable margin. Second pereopod (Figure 26A) overreaching antennal scale by less than length of fingers; fingers 1.9 to 2.6 times as long as palm; dactyl

(Figure 26f) armed with 17 to 26 spines on opposable margin. Third pereopod (Figure 26m) overreaching extreme anterior margin of carapace by

lengths of dactyl, propodus, and carpus; exopod

not nearly reaching distal end of ischium; ischium

and merus each bearing 2 strong spines near flexor

margins, carpus with 1; dactyl much shorter than

propodus. Fourth pereopod (Figure 26n) reaching

to about distal end of ischium of 2nd pereopod

when both extended anteriorly; dactyl longer

than propodus. Fifth pereopod (Figure 26o) similar to and more than $/4 as long as 4th, reaching

to about middle ys of ischium of 2nd pereopod

when both extended anteriorly; dactyl longer

than propodus.

Endopod of 1st pleopod of male (Figure 27a,b)

concave in distal portion of lateral margin, causFIGURE 27.—Leptochela (Leptochela) pugnax, male from

Nicobar Islands, carapace length 2.8 mm: a, right 1st pleopod; b, same, endopod; c, right 2nd pleopod; d, same,

appendix masculina and appendix interna. (Magnifications:

a, c, X 12; b, d, X 62.)

ing slight flaring of disolateral angle. Appendix

masculina (Figure 27c,d) bearing 6 long spines,

not overreaching appendix interna, not including

spines. Lateral branch of uropod (Figure 25e)

armed with 8 to 11 spines, in addition to setae.

SIZE.—Carapace lengths of males, 2.5-3.0 mm;

of nonbreeding females, 2.4-3.4 mm; of nonovigerous breeding females, 2.2-3.1 mm; of ovigerous females, 2.4-2.8 mm.

MATERIAL.—BAY OF BENGAL. Nicobar Islands; Investigator; received from Stanley W. Kemp: 11^ (2.5-2.8) 13$

(2.2-3.1) (3 ovig. (2.4-2.8)).

THAILAND. Anchorage in Mae Nam Chantaburi River

at Tha Chalaep Harbor, Chantaburi Province; 24 December

1957; George Vanderbilt Foundation Sta. 134: 1 $ (3.0) 1 9

(2.8).

VIET NAM. Bay of Nha Trang; 1 January to 10 April

1960; Ariel Gallardo St. 77: 1 $ (3.4).

PHILIPPINE ISLANDS. Cagmanaba Bay, Luzon (13°02'N,

123°17'E); coral and sand; 11 March 1909; 8:00 A.M.; dynamite (8 shots in 1-9 m); Albatross: 1$ (2.8). Mansalay, Mindoro (anchorage); electric light at surface over 16.5 m; 3

June 1908; 9:00 P.M.; Albatross: 2$ (2.8,2.9)2$ (2.8,2.9).

HABITAT.—Little is known of the ecological requirements of L. pugnax. All of the documented

specimens have been taken in inshore waters ranging in depth from 8 to 140 meters on both sand

and mud bottoms, and they have been attracted

to lights at the surface on a few occasions.

TytE-LocALiTY.—Five Indonesian localities between Java and the Moluccas; 9-55 meters.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—Leptochela (L.) pugnax is known

with reasonable certainty from the Maldive Islands eastward to eastern Indonesia and northward to the south coast of Honshu, Japan (see

discussion of distribution of L. (L.) robusta), in

depths of 8 to 55 meters. A few specimens from the

Red Sea that I have been able to examine through

the courtesy of L. B. Holthuis seem to differ

slightly from the typical form. Whether they, and

possibly those recorded as L. pugnax from the

Eastern Mediterranean by Holthuis and Gottlieb

(1958:22), represent a distinct taxon can be determined only by further study that is beyond the

scope of this review.

REMARKS.—Apparently only two described species of Leptochela, L. (L.) pugnax and the American L. (L.) serratorbita, invariably have the ventral angle of the orbit armed with a distinct tooth.

It seems probable, therefore, that the three young

specimens recorded by De Man (1920:30) as Leptochela sp. from as many additional Indonesian

localities are identifiable with L. (L.) pugnax, for

all of them are described as having the ventral

angle of the orbit dentate.

9. Leptochela (Leptochela) robusta Stimpson

FIGURE 28

Leptochela robusta Stimpson, 1860:43.—De Man, 1916:148

[part]; 1920:20 [part], pis. 3, 4 [except fig. 7x].

DIAGNOSIS.—Rostrum variable, dorsal margin

usually straight, concave, or sinuous. Carapace

wits 3 longitudinal dorsal ridges in breeding females only. Orbital margin entire, not serrate,

with mesially directed tooth on ventral portion;

suborbital angle unarmed. Fifth abdominal somite

entire, without dorsal elevations or posterior

tooth. Telson with 2 pairs of dorsolateral spines

in addition to anterior mesial pair; posterior margin without pair of minute mesial spines in addition to usual 5 pairs of prominent spines. Antennal scale about 2/s

 as long as carapace. First pereopod with 25 to 44 spines on opposable margin of

movable finger. Second pereopod with 28 to 47

spines on opposable margin of movable finger.

Third pereopod with exopod not nearly reaching

distal end of ischium. Endopod of 1st pleopod of

male not flared distolaterally. Appendix masculina, not including spines, distinctly overreaching

appendix interna. Maximum carapace length 6.4

mm.

DESCRIPTION.—Rostrum variable, dorsal margin

usually straight, concave, or sinuous, rarely overreaching eyes. Carapace without median dorsal

carina in males and nonbreeding females, tricarinate in breeding females. Orbital margin entire,

not minutely serrate; suborbital angle rounded,

but ventral margin of orbit armed with mesially

directed tooth visible only in dorsolateral view

(Figure 28a).

FIGURE 28.—Leptochela (Leptochela) robusta, male from

Albatross Sta. 5596, carapace length 6.0 mm: a, orbital

region, dorsolateral aspect; b, right 1st pleopod; c, same,

endopod; d, right 2nd pleopod; e, same, appendix masculina

and appendix interna. Male from same station, carapace

length 3.0 mm: /, right appendix masculina and appendix

interna. Male from Albatross Sta. 5159, carapace length

4.8 mm: g, left appendix masculina and appendix interna.

Male from Albatross Sta. 5169, carapace length 5.1 mm:

h, right appendix masculina and appendix interna. (Magnifications: b, d, X 6; a, X 12; c, X 25; e-h, X 62.)
NUMBER 222 35

Abdomen regularly rounded dorsally on 3 anterior somites and usually on 4th. Fifth somite

rounded or obscurely carinate, rarely distinctly

so; dorsal margin entire, without elevations of any

kind; posterior margin unarmed. Sixth somite

about 1.8 times as long as high, with usual tranverse swelling and carina near anterior end of dorsal surface, relatively short and obscure sharp

spine on ventrolateral surface, and distinct acute

tooth on posterodorsal margin of posterolateral

lobe. Telson, not including posterior spines, less

than H/2 times as long as 6th somite, about 3i/3

times as long as wide, armed with 2 pairs of dorsolateral. spines, spaced almost equidistantly, in addition to anterior mesial pair; posterior margin

without minute pair of spines between bases of

mesial pair of usual 5 pairs of prominent spines.

Eye with papilla on mesial surface of stalk proximal to cornea, cornea distinctly wider than stalk.

Antennular peduncle with stylocerite reaching

nearly as far as distolateral margin of basal segment; 2nd segment subequal in length to distal

segment in mesial aspect, no more than ]/2 as long

in dorsal aspect. Dorsolateral flagellum about 3i/3

times as long as carapace; ventromesial flagellum

fully H/3 times as long as carapace.

Antennal scale 0.5 to 0.7 as long as carapace,

2.8 to 4.2 times as long as wide, lateral margin

faintly sinuous, distal tooth obscurely separated

from mesial margin of blade. Distal segments of

antennal peduncle nearly 2/3 as wide as scale,

reaching to about midlength of scale. Flagellum

slightly more than 3 times as long as carapace.

Mouth parts typical of genus. Third maxilliped

not nearly reaching distal end of antennal scale,

distal segment about y4 as long as penultimate

segment.

First pereopod overreaching antennal scale by

less than length of fingers; fingers 1.5 to 1.7 times

as long as palm; dactyl armed with 25 to 44 spines

on opposable margin. Second pereopod overreaching antennal scale by less than length of fingers;

fingers 1.8 to 2.1 times as long as palm; dactyl

armed with 28 to 47 spines on opposable margin.

Third pereopod overreaching extreme anterior

margin of carapace by about length of dactyl; exopod falling far short of distal end of ischium;

ischium armed with row of 4 stout spines near

flexor margin; merus with 6 or 7 longer stout

spines near flexor margin; dactyl variable, usually

longer than propodus. Fourth pereopod reaching

nearly to distal end of ischium of 2nd pereopod

when both extended anteriorly; dactyl usually

longer, occasionally slightly shorter than propodus. Fifth pereopod similar to and about as long

as 4th, reaching at least to midlength of ischium

of 2nd pereopod when both extended anteriorly;

dactyl slightly shorter than propodus.

Endopod of 1st pleopod of male (Figure 28&,c)

subelliptical, little more than 214 times as long

as wide. Appendix masculina (Figure 2Sd—h)

bearing 8 to 11 long spines, longest arising from

proximal half of length, distinctly overreaching

appendix interna, not including spines. Lateral

branch of uropod armed with 9 to 13 movable

spines, in addition to setae.

SIZE.—Carapace lengths of males, 3.0-6.0 mm;

of nonbreeding females, 3.7-6.2 mm. De Man's

(1920) description of the Siboga material indicates that ovigerous females have a carapace

length of about 6.4 mm.

MATERIAL.—PHILIPPINE ISLANDS. Cebu; stomach of

cardinal fish, Archamia lineolata; 26 June 1970; Y Haneda

and F. Tsuji: 1 $ (4.4) 1 $ (4.5). Off Zamboanga, Mindanao;

6°54'0O"N, 122°04'30"E; surface over depth of 17 m; 10

October 1909; 6:00 P.M.; 10-foot plankton net (ship at

anchor); Albatross Sta. 5596: 2$ (3.0, 6.0) 4$ (3.7-6.2). Off

Tinagta Island, Tawitawi Group, Sulu Archipelago;

5°ll'50"N, 119°54'E; 18 m; coral sand; 21 February 1908;

10:08 A.M.; 9-foot Johnston oyster dredge; Albatross Sta.

5159: 13 (4.8). Off Sibutii Island. Sulu Archipelago;

4°32'15"N, 119°22'45"E; 18 m; coral sand; 27 February 1908;

8:36 A.M.; 9-foot Johnston oyster dredge; Albatross Sta. 5169:

1$ (5.1).

HABITAT.—Six of the 10 specimens examined

were taken at the surface over a depth of 17 meters. Two were found in the stomach of a cardinal

fish, and the remaining two were taken on coral

sand in 18 meters with a dredge modified from a

beam trawl by bolting a rake bar to the heel. The

lot of 61 Siboga specimens (De Man, 1920) that

almost certainly belong to this species were taken

on a bottom of fine and coarse sand with dead

shells in 141 meters.

TYPE-LOCALITY.—"Mari Sinensi, prof. 20 org."

It seems reasonably safe to assume that Stimpson's

"Mari Sinensi" refers to what is currently called

the South China Sea, because of references in

parts of his "Prodromus descriptionis" to "In mari

Sinensi Boreali, lat. bor. 23°." On the same page

with his preliminary description of L. (L.) ro-
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

busta, however, Stimpson records specimens of

Sicyonia ocellata "In mari Sinensi quoque, lat.

bor. 24°," a locality in Formosa Strait which connects the South China Sea and the East China

Sea, so a shadow of doubt must remain.

DISTRIBUTION.—Known with reasonable certainty only from the "China Sea"; off Cebu and

the Sulu Archipelago in the Philippine Islands;

and north of Pulau Waigeo, Indonesia; at the surface and to a depth of 141 meters.

Whether either L. (L.) robusta or L. (L.) irrobusta live in Japanese waters is uncertain at

the present time. The following passage is extracted from remarks kindly translated by Keiji

Baba from Kubo (1955:98):

Four species of this genus occur in Japan, including the two

reported here for the first time, but one of them, L. robusta,

is unavailable to me. Leplochela robusta was first reported

by Stimpson (1860) from the East China Sea and it has been

recorded subsequently from various localities in the Pacific

and Indian oceans. From Japanese waters, it was collected

from Toyama-wan by Kikuchi (1932) and from Tanabe-wan

(7-27 m), Matoya-wan (7 m), Gokasho-wan (13 m), Agowan (10-21 m), and Nanao-wan by Miyadi (1940, 1941). All

of Miyadi's specimens were identified by me at Prof. Miyadi's

request, but recently a question arose about that identification, and I again examined those from Tanabe-wan and

Ago-wan. This examination verified that those reported as

L. robusta are L. pugnax. Those from Matoya-wan, Gokashowan, and Nanao-wan have not been re-examined as yet. The

Toyama-wan record by Kikuchi was merely extracted from a

list, so it cannot be accepted as a reliable record of L.

robusta.

REMARKS.—Because of the brevity of Stimpson's

preliminary description, his failure to deposit any

material of the species in the British Museum,

and the subsequent loss of the invertebrates collected by the North Pacific Exploring Expedition

in the Chicago fire of 1871, the true identity of L.

(L.) robusta may never be known. We are inclined to believe, however, that it is identical

with the larger of the two forms so fully described

and figured by De Man (1920). Stimpson indicated that the species is one inch long, he stressed

its robust form, and he compared it with L. (L.)

gracilis, another of the larger species of the genus.

Such statements would hardly be applicable to

L. (L.) aculeocaudata, L. (L.) irrobusta, or L.

(L.) pugnax, the only other species known from

the region to which Stimpson's description might

otherwise apply.

There seemed to be no need to duplicate De

Man's excellent figures of this species. For a comparison of L. (L.) robusta and L. (L.) irrobusta,

see "Remarks" lnder the latter species.

10. Leptochela (Leptochela) serratorbita Bate

FICURES 29-31

Leptochela serratorbita Bate, 1888:859, pi. 139: fig. 1.—Young,

1900:497.—Rathbun, 1901:127.—Schmitt, 1935:134, fig. 7 —

Lunz, 1939:335.—Springer and Bullis, 1956:10.—Williams,

1965:41, figs. 33, 34.—Chace, 1972:16.

Leptochela serratorbita?.—Chace, 1937:111.

DIAGNOSIS.—Rostrum with dorsal margin

straight, concave, or sinuous, rarely convex. Carapace without longitudinal dorsal ridges in either

males or females. Orbital margin minutely spinulose dorsolaterally, with mesially directed rectangular or obtuse tooth on ventral portion; suborbital angle dentate. Fifth abdominal somite entire,

without dorsal elevations or posterior tooth. Telson with 2 pairs of dorsolateral spines in addition

to anterior mesial pair; posterior margin with pair

(frequently fused, rarely absent) of minute mesial

spines in addition to usual 5 pairs of prominent

spines. Antennal scale more than i/z but less than

y3 as long as carapace. First pereopod with 9 to 29

spines on opposable margin of movable finger.

Second pereopod with 13 to 37 spines on opposable margin of movable finger. Third pereopod

with exopod not nearly reaching distal end of

ischium. Endopod of 1st pleopod of male rather

narrowly rounded distally. Appendix masculina,

not including spines, occasionally reaching to or

slightly beyond distal end of appendix interna.

Maximum carapace length 4.1 mm.

DESCRIPTION.—Rostrum (Figure 29a,b) with

dorsal margin variable, straight, concave, sinuous,

or rarely convex, falling slightly short of or reaching slightly beyond level of distal margin of basal

segment of antennular peduncle. Carapace without median or dorsolateral carinae or ridges in

either males or females. Orbital margin (Figure

29b,c) minutely spinulose dorsolaterally, with

mesially directed rectangular or obtuse tooth on

ventral portion; suborbital angle dentate.

Abdomen (Figure 29rf) rounded dorsally on 3

anterior somites, usually bluntly carinate on at

least posterior part of 4th and all of 5th. Fifth
NUMBER 222 37

FIGURE 29.—Leptochela (Leptochela) serratorbita, ovigerous female from Key West, Florida,

carapace length 3.0 mm: a, carapace and anterior appendages; b, orbital region, lateral aspect;

c, anterior part of carapace and eyes, dorsal aspect; d, abdomen; e, posterior end of 6th

abdominal somite; /, telson and uropods; g, posterior margin of telson; h, same, mesial spines;

i, right antennule, dorsal aspect; /, right antenna, ventral aspect. (Magnifications: a, d, f,

X 12; b, c, e, i, j , X 25; g, X 62; h, X 260.)

somite entire, without dorsal prominences or posterodorsal tooth. Sixth somite nearly twice as

long as high, with usual transverse swelling near

anterior end of dorsal surface, relatively short

slender spine on ventrolateral surface, and rather

strong acute tooth on posterodorsal margin of posterolateral lobe (Figure 29<?). Telson (Figure

29/) nearly \\/2 times as long as 6th somite, fully

3 times as long as wide, armed with 2 pairs of

dorsolateral spines, posterior pair near midlength,

in addition to anterior mesial pair; posterior margin (Figure 29g) bearing pair (frequently fused,

rarely absent) of minute spines (Figure 29h) between bases of mesial pair of usual 5 pairs of

prominent spines.

Eye (Figure 29c) with papilla (visible only in

oblique view) on dorsomesial surface of stalk at

juncture with cornea, cornea slightly wider than

stalk.

Antennular peduncle (Figure 29i) with stylocerite reaching about as far as distolateral margin

of basal segment; 2nd segment about as long as

distal segment in ventromesial aspect, much

shorter in dorsal aspect.

Antennal scale (Figure 29;) 0.5 to 0.6 as long

as carapace, 2.9 to 4.4 times as long as wide, lateral margin noticeably concave at about midlength, distal tooth usually continuous with mesial

margin of blade, latter rarely with suggestion of

subdistal shoulder. Distal segments of antennal
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 30.—Leptochela (Leptochela) serratorbita, ovigerous female from Key West, Florida,

carapace length 3.0 mm: a, right mandible; b, right 1st maxilla; c, right 2nd maxilla; d, right

1st maxilliped; e, right 2nd maxilliped; /, right 3rd maxilliped; g, right 1st pereopod; h, same,

fingers; i, right 2nd pereopod; /, same, fingers; k, right 3rd pereopod; /, right 4th pereopod;

•m, right 5th pereopod. (Magnifications: c-g, i, k-m, X 25; a, b, h, j , X 62.)
NUMBER 222 39

peduncle about y4 as wide as scale, not nearly

reaching midlength of scale; ventral tooth on

basal segment neither prominent nor very sharp.

Mouthparts as illustrated (Figure 30a-/). Mandible (Figure 30a) with distal margin faintly

concave. Third maxilliped (Figure 30/) rarely

reaching as far as distal end of antennal scale, distal segment more than % a s

 l°ng

 a s

 penultimate

segment.

First pereopod (Figure 30g) usually overreaching antennal scale by less than length of fingers;

fingers 1.4 to 2.1 times as long as palm; dactyl

(Figure 30/J) armed with 9 to 29 spines on opposable margin. Second pereopod (Figure 30i) overreaching antennal scale by less than length of

fingers; fingers 1.5 to 2.4 times as long as palm;

dactyl (Figure 30;) armed with 13 to 37 spines on

opposable margin. Third pereopod (Figure 30A)

overreaching anterior margin of carapace by combined lengths of dactyl and propodus; exopod

not nearly reaching distal end of ischium; ischium

bearing row of 3 long spines near flexor margin

and subdistal spine on extensor margin; merus

armed with 5 or more similar spines near flexor

margin; dactyl distinctly shorter than propodus.

Fourtli pereopod (Figure 30/) reaching nearly to

midlength of ischium of 2nd pereopod when both

extended anteriorly; dactyl usually longer than

propodus. Fifth pereopod (Figure 30m) similar

to and little shorter than 4th, reaching to about

midlength of ischium of 3rd pereopod when both

extended anteriorly; dactyl usually longer than

propodus.

Endopod of 1st pleopod of male (Figures 31a,

b) rather narrowly rounded distally, lateral margin convex, not flared distally. Appendix masculina (Figure 31c,d) bearing 6 long spines and

occasionally reaching to end of or slightly beyond

appendix interna, not including spines. Lateral

branch of uropod (Figure 29/) armed with 7 to

13 spines.

SIZE.—Carapace length of males, 1.5-3.7 mm;

of females without eggs, 1.6-4.1 mm; of ovigerous

females, 2.7-3.9 mm.

MATERIAL.—NORTH CAROLINA. Beaufort; 6 November

1929; J. S. Gutsell: 1 9 (4.1).

GULF OF MEXICO: Cayos Areas, Bahi'a de Campeche;

18 m; II December 1952; from bonefish stomach; Oregon:

1$ (2.8).

FLORIDA KEYS. Key West; 1884; electric light at surface;

Albatross (W. Nye): 813 (2.5-3.4) 43$ (2.0-3.6) (4 ovig.

(2.9-3.0)).

PUERTO RICO. Playa de Fajardo; 23 February 1933;

8-foot circular net under cargo light; Johnson-Smithsonian

Deep-sea Expedition: 115 <J (2.0-3.7) 35$ (1.7-3.6) (1 ovig.

(3.4)) 1?. Cayo Icacos; 24 February 1933; 8-foot circular net

with cargo light and submarine light; Johnson-Smithsonian

Deep-sea Expedition: 3^ (1.8-2.6) 5$ (2.2-3.0). Off Isla de

Vieques; Culebritas lighthouse NE 1/2 N 18.5 km; 27 m;

coral; 8 February 1899; tangle; Fish Hawk Sta. 6091: 1 ovig.

2 (2.7). Off Isla de Culebra; Culebritas lighthouse NE 9.7

km; 27 m; coral; 8 February 1899; tangle; Fish Hawk Sta.

6093: 4 $ (2.5-3.0). Canal de Luis Pena, Isla de Culebra; 25

February 1938; 8-foot circular net under cargo light; JohnsonSmithsonian Deep-sea Expedition: 25 <J (2.1-3.3) 2$ (2.6,

2.8). Same; 3-foot net towed from port launch: 4 $ (2.3-2.5).

VIRGIN ISLANDS. Brewers Bay, Saint Thomas; 1 March

1933; 8-foot circular net under cargo light; Johnson-Smithsonian Deep-sea Expedition: 15 $ (1.5-2.4) 53$ (1.6-2.8).

Saint Thomas; 17-24 January 1884; Albatross: 83 (1.9-3.0)

2$ (2.6) 1? (2.3). Sopers Hole, Tortola; 5-11 m; shells and

compact sand; 1 April 1958; dredge; Smithsonian-Bredin

Caribbean Expeditions Sta. 25-58; 2 $ (2.6, 2.7). Off Vixen

Point, Prickly Pear Island, Virgin Gorda; 6 April 1958; 9:00

P.M.; light at bottom at anchorage in 20 m; SmithsonianBredin Caribbean Expeditions Sta. 36b-58: 1 juv. (0.8).

LEEWARD ISLANDS. Off Oyster Pond Landing, west side

of Barbuda; 5 April 1956; 7:00-8:30 P.M.; light over side at

anchorage in 5i/£ m; Smithsonian-Bredin Caribbean Expeditions Sta. 84-56: 222 $ (2.3-3.6) 200$ (2.6-4.0) (8 ovig.

(3.0-3.9)). South of Bird Island, Nonsuch Bay, Antigua Island; 23 April 1958; light over side at anchorage in 7 m;

Smithsonian-Bredin Caribbean Expeditions Sta. 93-58: 1 $

(3.4).

a

FIGURE 31.—Leptochela (Leptochela) serratorbita, male from

Key West, Florida, carapace length 3.1 mm: a, right 1st

pleopod; b, same, endopod; c, right 2nd pleopod; d, same,

appendix masculina and appendix interna. (Magnifications:

a, c, X 12; b, d, X 62.)
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HABITAT.—In or over depths of 514 to 27 meters.

It may be noted that 8 of the 15 lots and all but

25 of the 826 specimens studied were taken under

a light at the surface at night. See discussion of

the habitat of L. (L.) bermudensis and remarks

under L. (P.) carinata.

TYPE-LOCALITY.—Saint Thomas, Virgin Islands.

DISTRIBUTION.—North Carolina and western

Gulf of Mexico to the Leeward Islands, in and

over depths of 5\A to 27 meters.

Reexamination of the material collected at the

surface beneath a light and dredged in 15-29

meters on a sandy bottom at San Lucas, Baja

California, tentatively assigned to L. (L.) serratorbita by Chace (1937) has revealed no obvious

diflerences from western Atlantic populations,

but a more thorough comparison would be desirable when additional adult specimens become

available from the eastern Pacific.

REMARKS.—The available evidence would seem

to indicate that L. (L.) serratorbita has a more

restricted range in the western Atlantic, both geographically and bathymetrically, than L. (L.) bermudensis, but that it is by far the commonest species of Leptochela within the limits of its range.

11. Leptochela (Leptochela) sydniensis

Dakin and Colef ax

FICURES 32-34

Leptochela aculeocaudata.—Kemp, 1915:311 [part], figs. 34,

35, pi. 13: fig. 14; 1925:254.—Menon, 1937:6, pis. 1, 2: figs.

15-45.—Kubo, 1955:103, fig. 6.—Pillai, 1955:48, fig. 1.—

Fujino and Miyake, 1970:239. [Not L. aculeocaudata Paulson, 1875.]

?Leptochela hainanensis Yu, 1936:87, figs. 1-3.

Leptochela sydniensis Dakin and Colefax, 1940:153, figs. 245,

246.

DIAGNOSIS.—Rostrum with dorsal margin nearly

straight or sinuous. Carapace with 3 longitudinal

dorsal ridges in breeding females only. Orbital

margin entire, not serrate, without mesially directed tooth on ventral portion; suborbital angle

unarmed. Fifth abdominal somite entire, without

dorsal elevations or posterior tooth. Telson with

1 pair of dorsolateral spines in addition to anterior

mesial pair; posterior margin normally without

pair of minute mesial spines in addition to usual

5 pairs of prominent spines. Antennal scale about

i/2 as long as carapace. First pereopod with 20 to

44 spines on opposable margin of movable finger.

Second pereopod with 21 to 46 spines on opposable margin of movable finger. Third pereopod

with exopod nearly reaching distal end of ischium. Endopod of 1st pleopod of male obliquely

rounded distally, lateral margin slightly concave

but not flared distally. Appendix masculina, not

including spines, usually not overreaching appendix interna. Maximum carapace length 4.7 mm.

DESCRIPTION.—Rostrum (Figure 32a) with dorsal margin nearly straight or sinuous, sometimes

overreaching eyes but seldom reaching level of

distal margin of basal segment of antennular peduncle. Carapace with median dorsal carina on

anterior i/3 or less of length in males and nonbreeding females, extending to midlength or beyond and with parallel but usually indistinct dorsolateral ridges in breeding females; ridges low, not

concealing dorsal midline from lateral view; anterior lobe of carapace not concealing basal segment of antennal peduncle from lateral view.

Orbital margin (Figure 326) entire, not spinulose, without distinct mesially directed tooth on

ventral portion; suborbital angle rounded, unarmed.

Abdomen (Figure 32c) rounded dorsally on 3

anterior somites, often partially carinate on 4th,

usually distinctly carinate on 5th. Fifth somite

entire, without dorsal prominences or posterodorsal tooth. Sixth somite nearly twice as long as

high, with transverse carinate swelling at anterior

end; usual slightly curved spine on ventrolateral

surface and distinct acute tooth on posterodorsal

margin of posterolateral lobe (Figure 32d). Telson (Figure S2e) nearly \i/2 times as long as 6th

somite, about 3y2 times as long as wide, armed

with 1 pair of dorsolateral spines at about midlength, in addition to anterior mesial pair; posterior margin (Figure 32/) normally without pair

of minute spines between bases of mesial pair of

usual 5 pairs of prominent spines.

Eye (Figure 326) with rather prominent papilla (best seen in dorsolateral view) on dorsomesial surface of stalk at jucture with cornea, cornea inflated, wider than stalk.

Antennular peduncle (Figure 32g) with stylocerite reaching nearly as far as distolateral margin of basal segments; 2nd segment nearly as long

as distal segment in both mesial and dorsal aspects.
NUMBER 222 41

FIGURE 32.—Leptochela (Leptochela) sydniensis, ovigerous female from Port Arthur, Tasmania,

carapace length 4.4 mm: a, carapace and anterior appendages; b, anterior part of carapace and

eyes, dorsal aspect; c, abdomen; d, posterior end of 6th abdominal somite; e, telson and

uropods; /, posterior margin of telson; g, left antennule, dorsal aspect; h, left antenna, ventral

aspect. (Magnifications: a, c, e, X 12; b, d, g, h, X 25; /, X 62.)
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 33.—Leptochela (Leptochela) sydniensis, ovigerous female from Port Arthur, Tasmania,

carapace length 4.4 mm: a, right mandible; b, right 1st maxilla; c, right 2nd maxilla; d, right

1st maxilliped; e, left 2nd maxilliped; /, right 3rd maxilliped; g, left 1st pereopod; h, same,

fingers; i, left 2nd pereopod; /, same, fingers; k, left 3rd pereopod; /, right 4th pereopod;

m, right 5th pereopod. (Magnifications: /, g, i, k-m, X 12; c-e, X 25; a, b, h, j , X 62.)
NUMBER 222 43

Antennal scale (Figure 32h) about \/2 as long

as carapace, 3.1 to 3.9 times as long as wide, lateral

margin distinctly concave near midlength, junction of mesial margin of blade with distal tooth

marked only by faint sinuosity, without distinct

shoulder. Distal segments of antennal peduncle

nearly 2/s as wide as scale, extending to about

midlength of scale; basal segment bluntly produced distoventrally, without acute tooth.

Mouthparts as illustrated (Figure 33a-/). Third

maxilliped (Figure 33/) rarely reaching level of

tip of antennal scale, distal segment about y4 as

long as penultimate segment.

First pereopod (Figure 33g) occasionally overreaching antennal scale by as much as length

of fingers; fingers 1.3 to 1.9 times as long as palm;

dactyl (Figure SSh) armed with 20 to 44 spines

on opposable margin. Second pereopod (Figure

33J) occasionally overreaching antennal scale by

more than length of fingers; fingers 1.4 to 2.3

times as long as palm; dactyl (Figure 33;) armed

with 21 to 46 spines on opposable margin. Third

pereopod (Figure 33ft) overreaching extreme

anterior margin of carapace by about length of

dactyl; exopod not nearly reaching distal end of

ischium; ischium armed with 3 spines on flexor

margin; merus with 4 spines; carpus with 1 spine;

dactyl distinctly shorter than propodus. Fourth

pereopod (Figure 33/) reaching distal end of ischium of 2nd pereopod when both extended anteriorly; dactyl usually distinctly longer than propodus. Fifth pereopod (Figure 33m) similar and

subequal in length of 4th, barely overreaching

midlength of ischium of 2nd pereopod when both

extended anteriorly; dactyl about as long as propodus.

Endopod of 1st pleopod of male (Figure 34a,6)

obliquely rounded distally, lateral margin slightly

concave but not flared distally. Appendix masculina (Figure 34c,d) bearing 5 to 7 long spines,

usually not overreaching appendix interna, not

including spines. Lateral branch of uropod (Figure

$2e) armed with 10 to 15 movable spines.

SIZE.—Carapace lengths of males, 2.3-4.4 mm;

of females without eggs, 2.3-4.7 mm; of ovigerous

females, 3.3-4.4 mm. The smallest male, from the

East China Sea, has the appendix masculina no

more than i/2 as long as the appendix interna.

MATERIAL.—EAST CHINA SEA. Northeast of Taiwan;

d

FIGURE 34.—Leptochela (Leptochela) sydniensis, male from

north of Cape Howe, Victoria, Australia, carapace length

4.4 mm: a, left 1st pleopod; b, same, endopod; c, left 2nd

pleopod; d, same, appendix masculina and appendix interna.

(Magnifications: a, c, X 12; b, d, X 62.)

26o05'N, 122°55'E; 110-111 m; Kaiyo-Maru Sta. 9: 3,$ (2.3-

3.4) 79 (35-4.1) (2 ovig. (3.5, 4.1)) (Zoological Laboratory

of Kyushu University).

NEW SOUTH WALES. North Head, Sydney; 66 m; 27

February 1973; dredge; Australian Museum Shelf Benthic

Survey Sta. 35: 3$ (35-4.3) (1 ovig. (?)) (Australian Museum). 1.6 km E of Malabar outlet, Sydney; 33°58'15"S,

150°17'E; 66 m; 26 June 1973; dredge; AMSBS Sta. 3: 10 3

(3.1-3.7) 25 9 (3.4-4.3) (8 ovig. (3.3-3.6)) (Aus. Mus.). E of

Malabar, Sydney; 83 m; 29 May 1974; AMSBS Sta. 40: 3$

(3.7-4.0) (Aus. Mus.). 2 km E of Long Bay; 33°58'50"S,

150°17'E; 66 m; 30 January 1974; dredge; AMSBS Sta.

4: 20 $ (2.8-45) 8$ (3.6-4.4) (1 ovig. (?)) (Aus. Mus.). 2.3

km E of Malabar, Sydney; 33°59^7"S, 15°16'48"E; 66 m; 29

January 1974; dredge; AMSBS Sta. 5: 5$ (3.8-4.0) 3$ (3.8-

4.2) (Aus. Mus.).

VICTORIA. N of Cape Howe; 37°26'S, 150°15'E; 149 m;

19 June 1962; CSIRO Fisheries Sta. G2/59/62: 1$ (4.4)

(Aus. Mus.). N of Cape Howe; 37°26.5'S, 150°l7'E; 300 m;

19 June 1962; CSIRO Fisheries Sta. G2/60/62: 1 $ (3.4) 1 $

(4.7) (Aus. Mus.). W of Cape Everard; 37°55'S, 149°00'E;

77 m; 20 June 1962; CSIRO Fisheries Sta. G2/61/62: 2$

(2.2+, 4.2) (Aus. Mus.).

SOUTH AUSTRALIA. S of Investigator Group; 34°S,

134°32'E; 152 m; 12 July 1962; CSIRO Fisheries Sta. G2/

128/62: 13 (4.4) 2 9 (3.2, 3.6) (1 ovig. (3.6)) (Aus. Mus.).

WESTERN AUSTRALIA. Great Australian Bight; 35°43'S,

125°04'E; 80 m; 7 July 1962; CSIRO Fisheries Sta. G2/104/62:

\$ (3.4) (Aus. Mus.).
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TASMANIA. In Port Arthur; C. T. Harrison: 1 ovig. $

(4.4) (Aus. Mus.).

HABITAT.—Kemp (1915) recorded specimens

from weeds in shallow water on the extreme south

coast of India, but most of the other records here

attributed to this species pertain to depths greater

than two meters. Fujino and Miyake (1970) listed

material from the East China and Yellow seas

from depths of 28 to 110 meters, as well as from

the surface. The Australian specimens recorded

above were taken in depths of 66 to 300 meters.

Inasmuch as the larger lots from off New South

Wales were taken with a dredge in 66 meters, it

seems likely that the species lives on or near the

bottom at least part of the time. The surface records from the East China or Yellow seas, on the

other hand, suggest that it is also pelagic at times.

TYPE-LOCALITY.—Off Sydney, New South Wales,

Australia. No type-material of this species could

be found in the Australian Museum or in the University of Sydney.

DISTRIBUTION.—If my conclusions are correct

(see "Remarks"), L. sydniensis is known from the

Arabian Sea, the Bay of Bengal, the South China

and Yellow seas, and Japanese waters, as well

as off the southeastern and southern coasts of

Australia.

REMARKS.—The opportunity to examine specimens of L. aculeocaudata from the Red Sea has

convinced me that the concept of that species

adopted by Kemp (1915, 1925) and subsequent

authors is probably incorrect. Kemp (1925:250)

was the first to discover that the presence or absence of paired dorsolateral longitudinal ridges

on the carapace was not a simple sexual character

but that these ridges are developed only in breeding females of certain species of the genus. He

concluded his remarks on the subject as follows:

"Dimorphic forms of the female have been found

in L. robusta, L. aculeocaudata and L. pugnax,

and it is not unlikely that the phenomenon occurs

throughout the genus." We now know that the

so-called tricarinate carapace is restricted to

breeding females in most, but not all, of the species; it is found in all specimens of both sexes of

L. (L.) aculeocaudata and L. (L.) japonica and

in no specimens of either sex of L. (L.) gracilis

and L. (L) serratorbita. What, then, is the identity of the species that Kemp assigned to L.

aculeocaudata?

I at first believed that L. hainanensis Yu, 1936,

was the earliest available name for this form. Yu's

species seems to agree with the one described by

Kemp in most characters, as pointed out by Armstrong (1941) and Fujino and Miyake (1970),

and its type-locality lies between the populations

in the Indian Ocean recorded by Kemp and those

in the western North Pacific recorded by Kubo

(1955) and Fujion and Miyake. Reexamination

of material from Hainan may reveal that this is

the correct solution, but I am sufficiently disturbed

by the figure of the antennal scale given by Yu

(1936, fig. 1) and by his statement that "the outer

margin is nearly straight, not concave behind the

middle" to resist adopting the name L. hainanensis for this species at this time. According to Yu's

figure, the antennal scale not only lacks the wellmarked concavity in the lateral margin, but the

blade seems to form a rather distinct shoulder at

the base of the distal spine.

In describing L. sydniensis, Dakin and Colefax

(1940:156) noted that the species was allied to

L. robusta, L. aculeocaudata, and L. pugnax, according to the key published by Kemp (1925:

250). They proceeded to note the differences between the Australian species and L. robusta and

L. pugnax, but they failed to distinguish it from

Kemp's interpretation of L. aculeocaudata. The

suspicion that there might be no differences has

been supported by the opportunity afforded

through the kind cooperation of Dr. Fujino to

examine part of the lot from the southernmost

station in the East China Sea recorded by Fujino

and Miyake as L. aculeocaudata. No obviously

significant differences could be found between

these specimens and those from Australia. In the

three females with a tricarinate carapace from the

East China Sea, the dorsolateral ridges are stronger

and sharper than in those from Australia, but the

importance of this apparent distinction could not

be evaluated from the material available to me.

It is very possible that this interpretation does

not represent the final taxonomic word on this

rather complex problem, but I hope that I may

have contributed to some extent toward such a

solution.

Proboloura, new subgenus

DEFINITION.—Sixth abdominal somite bearing
NUMBER 222 45

movable lappet or flap attached anteriorly to

transverse carinate swelling on dorsal surface. Telson with anterior pair of dorsolateral spines nearly

in line with anterior dorsal pair. Antennal scale

usually at least y4 as long as carapace. Third

pereopod with exopod reaching nearly or quite

as far as distal end of ischium. Fifth pair of pereopods reduced, about 2/3 as long as 4th pair.

TYPE-SPECIES.—Leptochela carinata Ortmann.

ETYMOLOGY.—From probolos (G., = projecting

object or prominence) + oura (G., = tail), in

reference to lappet on 6th abdominal somite.

Gender: Feminine.

REMARKS.—Leptochela carinata is so different

from the other members of the genus, especially

in the presence of a dorsal lappet on the sixth

abdominal somite and the disposition of the dorsolateral spines on the telson, that supraspecific distinction seems desirable. That full generic status

is hardly justified at this time is indicated by the

much greater similarity between L. carinata and

the other species of the genus than between any

species of Leptochela and the species of other

pasiphaeid genera. Also, the gap between L. carinata and its congeners is partially filled by L.

japonica Hayashi and Miyake, in which the fifth

abdominal somite is similarly crested and the fifth

pereopods are somewhat reduced.

12. Leptochela (Proboloura) carinata Ortmann

FIGURES 35-37

Leptochela carinata Ortmann, 1893:41 [part], pi. 4: fig. 1.—

Rathbun, 1901:127.—Schmitt, 1935:134.—Hayashi and

Miyake, 1969:6, fig. 3.—Chace, 1972:16.

DIAGNOSIS.—Rostrum with dorsal margin usually convex, occasionally straight or even concave.

FIGURE 35.—Leptochela (Proboloura) carinata, ovigerous female from Grampus Sta. 5084,

carapace length 6.7 mm: a, carapace and anterior appendages; b, rostrum and orbit, dorsolateral aspect; c, abdomen; d, anterior end of 6th abdominal somite, dorsal aspect showing

lappet; e, posterior end of 6th abdominal somite. Male from Rum Cay, Bahamas, carapace

length 5.7 mm: /, telson and uropods; g, posterior margin of telson; h, same, mesial part.

(Magnifications: a, c ,X 6; b, d-f, X 12; g, X 25; h, X 260.)
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Carapace with 3 longitudinal dorsal ridges in

breeding females only. Orbital margin entire, not

serrate, with mesially directed tooth on ventral

margin; suborbital angle unarmed. Fifth abdominal somite with dorsal carina elevated into 4 teeth,

anterior 2 very prominent, posteriormost forming

blunt projection on posterior margin. Telson

with 2 pairs of dorsolateral spines in addition to

anterior mesial pair; posterior margin without

pair of minute mesial spines in addition to usual

5 pairs of prominent spines. Antennal scale usually more than y4 as long as carapace. First pereopod with 41 to 58 spines on opposable margin of

movable finger. Third pereopod with exopod

reaching nearly or quite as far as distal end of

ischium. Endopod of 1st pleopod of male rounded

distally, not produced distolaterally. Appendix

masculina, not including spines, not overreaching

appendix interna. Maximum carapace length 7.7

mm.

DESCRIPTION.—Rostrum (Figure 35a) with dorsal margin usually somewhat convex, occasionally

straight or even concave, short, not overreaching

basal segment of antennular peduncle. Carapace

with median dorsal carina on anterior y3 to i/2

of length but without paired dorsolateral carinae

in males and nonbreeding females, dorsally tricarinate over most of length in breeding females.

Orbital margin (Figure 35&) entire, not spinulose, but with mesially directed tooth on ventral

portion; suborbital angle rounded.

Adbomen (Figure 35c) regularly rounded dorsally on 3 anterior somites and at least all but

posterior i/3 of 4th. Fifth somite sharply carinate,

carina produced into 2 prominent acute anterior

teeth and 2 low blunt teeth, posteriormost on posterior margin of somite. Sixth somite about li/£

times as long as high, with movable lappet subtriangular (broadening distally) (Figure 35d),

comparatively short spine (Figure 35e) on ventrolateral margin, and acute, sometimes rather obscure tooth on posterodorsal margin of posterolateral lobe. Telson (Figure 35/), not including

posterior spines, about 1.6 times as long as 6th

somite, less than 3 times as long as wide, margins

noticeably sinuous, armed with 3 pairs of long

spines, 1 pair anteromesial, 2 pairs dorsolateral,

anterior pair of latter nearly in line with anteromesial pair, posterior pair arising at about midlength of telson; posterior margin (Figure 35g)

with indistinct lobe (Figure $5h) but without

pair of minute spines between bases of mesial

pair of usual 5 pairs of prominent spines.

Eye with papilla on mesial surface of stalk proximal to cornea, cornea distinctly wider than stalk.

Antennular peduncle (Figure 36a) with stylocerite considerably overreaching basal segment;

2nd segment subequal to distal segment in mesial

aspect but distinctly shorter in dorsal aspect.

Antennal scale (Figure 366) 0.7 to 0.9 times as

long as carapace, 4.0 to 5.0 times as long as wide,

both lateral and mesial margins nearly straight,

latter continuous with distal spine without any

indication of subdistal shoulder. Distal segments

of antennal peduncle little more than 14 as wide

as scale, reaching to about middle 1/3 of scale;

basal segment with prominent ventral spine.

Mouthparts as illustrated (Figures $6c-h).

Third maxilliped (Figure 36/i) overreaching antennal scale, distal segment about 2/s

 as long as

penultimate segment.

First pereopod (Figure 36i) overreaching antennal scale by slightly more or, usually, less than

length of fingers; fingers 1.0 to 1.2 times as long

as palm; dactyl (Figure 36;) armed with 41 to 58

spines on opposable margin. Second pereopod

(Figure 36A) rarely overreaching antennal scale

by more than length of fingers; fingers 1.1 to 1.3

times as long as palm; dactyl (Figure 36/) armed

with 43 to 66 spines on opposable margin. Third

pereopod (Figure 36m) overreaching extreme

anterior margin of carapace by length of dactyl

and about i/2 of propodus; exopod reaching nearly

or quite as far as distal end of ischium; ischium

armed with row of about 4 spines near extensor

margin and about 6 on flexor margin; merus with

5 spines near extensor margin and 9 to 11 on flexor

margin; dactyl shorter than propodus. Fourth

pereopod (Figure 36n) reaching to middle i/3 of

ischium of 2nd pereopod when both are extended

anteriorly; ischium armed with several stout

spines on semicircular anterior lobe; similar

spines on flexor margins of merus, carpus, and

propodus; dactyl shorter than propodus. Fifth

pereopod (Figure 36o) reduced, not more than

y3 as long as 4 th pereopod, reaching barely to

distal end of basis of 2nd pereopod when both are

extended anteriorly, ischium and merus especially

short; ischium armed distally with several stout

spines; similar spines on flexor margins of merus,
NUMBER 222 47

FIGURE 36.—Leptochela (Proboloura) carinata, female from Grampus Sta. 5084, carapace length

7.7 mm: a, right antennule, dorsal aspect; b, right antenna, ventral aspect; c, right mandible;

d, right 1st maxilla; e, right 2nd maxilla; /, right 1st maxilliped; g, right 2nd maxilliped;

h, right 3rd maxilliped; i, right 1st pereopod; ;', same, fingers; k, right 2nd pereopod; /, same,

fingers. Male from Rum Cay, Bahamas, carapace length 5.7 mm: m, left 3rd pereopod; n, right

4th pereopod; o, right 5th pereopod. (Magnifications: a, b, h, i, k, X 6; e-g, tn-o, X 12;

carpus, and propodus; dactyl very small, considerably shorter than propodus.

Endopod of 1st pleopod of male (Figure $7a,b)

truncately rounded distally,, lateral margin not

flared distally. Appendix masculina (Figure $7c,d)

bearing 7 long spines, not overreaching appendix

interna, not including spines. Lateral branch of

uropod (Figure 35/) armed with 14 to 22 spines.

SIZE.—Carapace lengths of males, 3.0-5.7 mm;

of nonbreeding females, 2.4-7.7 mm; of ovigerous

females, 4.3 and 6.7 mm.

MATERIAL.—GEORGES BANK. 41°22'N, 68°18'W, 35 m;

9 December 1955; 1-meter conical stramin net mounted on

steel sled hauled along bottom; Albatross III Cruise 70, Collection 17 (R. L. Wigley): I $ (3.0) Northeast Fisheries Center, Woods Hole, Massachusetts).

GULF OF MEXICO. Between Cape San Bias and Tampa

Bay, Florida; 28°47'N, 83°42'W; 24 m; 1 February 1970;

dredge; L. G. Abele: \$ (5.7). Same; 28°31'N, 84°16'W;

26 m; corraline rubble; 9 April 1970; Peterson grab; L. G.

Abele: 1 9 (5.2). Anclote Light, Florida, E 7/8 N, 39.8 km;

23 m; rock, coral, and sand; 17.2°C; 28 March 1901 (10:30

A.M.);'dredge; Fish Hawk Sta. 7106: 2$ (4.4, 6.2). Between

Marco and Dry Tortugas, Florida; 25°44'32"N, 82O37'15"W;
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 37.—Leptochela (Proboloura) carinata, male from

northeastern Gulf of Mexico, carapace length 5.7 mm:

a, right 1st pleopod; b, same, endopod; c, right 2nd pleopod;

d, same, appendix masculina and appendix interim. (Magnifications: a, c, X 6; b, X 25; d, X 62.)

35 m; sand and coralline algae; 10 March 1889; hand dredge;

Grampus Sta. 5084: 2$ (6.7, 7.7) (1 ovig. (6.7)).

BAHAMAS. Rum Cay; surface; 1886; Albatross: 2$ (5.2,

5.7) 2 $ (5.3, 5.5).

PUERTO RICO. Off Isla de Vieques; San Juan lighthouse

NW 1/8 N 27.3 km; 20 m; coral, sand, and shells; 8 February 1899; tangle; Fish Hawk Sta. 6084: 1 $ (3.2). Same; Point

Mula lighthouse SSW 3/8 W 10.6 km; 26 m; coral and sand;

8 February 1899; 7-foot beam trawl; Fish Hawk Sta. 6085:

1£ (4.0). Same; Point Mula lighthouse E 1/2 N 20.8 km; 11

m; coral; 8 February 1899; tangle; Fish Hawk Sta. 6096: 1

ovig. $ (4.3). Off Isla de Culebra; Culebritas lighthouse NE

9.7 km; 27 m; coral; 8 February 1899; tangle; Fish Hawk Sta.

6093: 1 $ (4.3). Same; Canal de Luis Pena; 25 February 1933;

8-foot circular net under cargo light; Johnson-Smithsonian

Deep-sea Expedition: 1 9 (2.4).

HABITAT.—In depths of 11 to 35 meters on bottoms of coral and sand, as well as coralline algae,

rocks, and shells. Although 11 of the 16 specimens

examined were apparently taken on or near the

bottom with dredges, a beam trawl, a plankton

net, a Peterson grab, and tangles, the lot of four

specimens from the Bahamas was collected at the

surface, as was the single specimen taken by the

Johnson-Smithsonian Expedition in Canal de

Luis Pena, Isla de Culebra. Also included among

the material collected under a light at the latter

station were three L. (L.) bermudensis and 27

L. (L.) serratorbita, so it is possible that L. (P.)

carinata, which was taken at three bottom stations

in that general area, is not especially photopositive.

TYPE-LOCALITY.—Off Bafa de Maraj6 ("Kiistenbank vor der Tocantins-Miindung"), Estado

do Para, Brazil, 50-100 meters.

DISTRIBUTION.—Georges Bank; Gulf of Mexico;

Bahamas to Estado do Para, Brazil, in depths of

11 to more than 50 meters.

REMARKS.—See "Remarks" under L. (L.) bermudensis.

Although L. (P.) carinata has the most extensive known distribution of any of the four Atlantic species of the genus, it is less well represented

in the collections available to me than any of the

other three species. On the other hand, it is the

only one of the four species that was taken one

or more times with each of the other species:

once with L. (L.) papulata, once with L. (L.)

serratorbita, and once with both L. (L.) bermudensis and L. (L.) serratorbita. The relative paucity of L. (P.) carinata in collections may reflect

the possibility, mentioned above in the habitat

discussion, that the species is less attracted to light

than are either L. (L.) bermudensis or L. (L.)

serratorbita. An alternative explanation may be

that L. (P.) carinata does not swarm as do the

two latter species.
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