New and Rare Stomatopod Crustacea

from the

Indo-West-Pacific Region

RAYMOND B. MANNING

'

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 264
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SMITHSONIA N CONTRIBUTION S T O Z O O L O G Y • N U M B E R 26 4

New and Rare Stomatopod Crustacea

from the

Indo-West-Pacific Region

Raymond B. Manning

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978
ABSTRAC T

Manning, Raymond B. New and Rare Stomatopod Crustacea from the IndoWest-Pacific Region. Smithsonian Contributions to Zoology, number 264, 36

pages, 16 figures, 1978.—Nineteen species of stomatopod crustaceans are reported.

Five new genera are recognized: Hoplosquilloides, for H. coronatus, new species:

Keppelius, for Lysiosquilla hystricotelson Barnard, 1958; Pullosquilla, for Austrosquilla litoralis Michel and Manning, 1971, A. malayensis Manning, 1968,

and a new species, P. thomassini; Busquilla, for Squilla quadraticauda Fukuda,

1911, and B. plantei, new species; and Tuleariosquilla, for T. parvula, new

species. Five other new species are described: Alima orientalis, Clorida japonica,

Cloridopsis aquilonaris, Harpiosquilla malagasiensis, and Neocoronida martensi.

Four species are redescribed: Clorida minor (Jurich, 1904); Gonodactylus viridis

Serene, 1954; Neocoronida trachurus (von Martens, 1881); and Parvisquilla

multituberculata (Borradaile, 1898). Gonodactylus confinis de Man, 1902, and

G. segregatus Lanchester, 1903 are shown to be synonyms of G. affinis de Man,

1902; Coronis spinosa Wood-Mason, 1875 is a synonym of Coronis tricarinata

Claus, 1871; and Parvisquilla xishaensis Liu, 1975 is a synonym of Parvisquilla

multituberculata (Borradaile, 1898). Additional material of Clorida latispina

Manning, 1968 and C. malaccensis Manning, 1968 is reported; C. malaccensis

var. moluccensis Moosa, 1973 is a synonym of the latter.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Instftmion's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral

Montastrea cavemosa (Linnaeus).

Library of Congress Cauloging in Publication Data

Manning, Raymond B 1934-

New and rare stomatopod Crustacea from the Indo-West-Pacific region.

(Smithsonian contributions to zoology ; 264)

Bibliography: p.

1. Stomatopoda. 2. Rare animals—Indian Ocean. 3. Rare animals—Pacific Ocean. 4. Crustacea—Classification. 5. Crustacea—Indian Ocean. 6. Crustacea—Pacific Ocean. I. Title. II.

Series: Smithsonian Institution. Smithsonian contributions to zoology ; 264.

QL1.S54 no. 264 [QL444.M.375] 591'.08s [595'.382'09165] 77-14228
Contents

Page

Introduction 1

Acknowledgments 1

Family GONODACTYLIDAE Giesbrecht, 1910 2

Genus Gonodactylus Berthold, 1827 2

Gonodactylus affinis de Man, 1902 2

Gonodactylus viridis Serene, 1954 4

Hoplosquilloides, new genus 6

Hoplosquilloides coronatus, new species 6

Family LYSIOSQUILLIDAE Giesbrecht, 1910 8

Genus Heterosquilla Manning, 1963 8

Heterosquilla tricarinata (Claus, 1871) 8

Keppelius, new genus 9

Keppelius hystricotelson (Barnard, 1958) 10

Genus Neocoronida Manning, 1976 11

Key to Species of Neocoronida 11

Neocoronida martensi, new species 12

Neocoronida trachurus (von Martens, 1881) 13

Genus Parvisquilla Manning, 1973 15

Key to Species of Parvisquilla 16

Parvisquilla multituberculata (Borradaile, 1898) 16

Pullosquilla, new genus 18

Key to Species of Pullosquilla 19

Pullosquilla litoralis (Michel and Manning, 1971) 19

Pullosquilla thomassini, new species 20

Family SQUILLIDAE Latreille, 1803 21

Genus Alima Leach, 1817 21

Alima orientalis, new species 21

Busquilla, new genus 23

Busquilla plantei, new species 23

Genus Clorida Eydoux and Souleyet, 1842 25

Clorida japonica, new species 25

Clorida latispina Manning, 1968 26

Clorida malaccensis Manning, 1968 26

Clorida minor (Jurich, 1904) 27

Genus Cloridopsis Manning, 1968 28

Cloridopsis aquilonaris, new species 28

Genus Harpiosquilla Holthius, 1964 30

Harpiosquilla malagasiensis, new species 30

Tuleariosquilla, new genus 30

Tuleariosquilla parvula, new species 32

Literature Cited 34

New and Rare Stomatopod Crustacea

from the

Indo-West-Pacific Region

Raymond B. Manning

Introduction

Visits to European museums in 1971 and to the

Zoological Survey of India, Calcutta in 1972, and

examination of collections received since then have

resulted in the accumulation of a large amount of

new information on Indo-West-Pacific stomatopods.

Because some names are required by colleagues

working in other fields, redescription of five known

species, observations on types of several poorly

known species, and descriptions of new taxa, including five new genera and nine new species, are

presented here. More cohesive summaries of other

Indo-West-Pacific stomatopods, at the generic or

species-group levels are in preparation or in press;

these include reviews of Lysiosquilla and the members of the falcatus group of Gonodactylus and

redescriptions of types and descriptions of new species in Oratosquilla. A general review of stomatopod taxa at and above the generic level is also in

preparation.

All measurements are in millimeters (mm). TL

refers to total length, measured on the midline

from the anterior margin of the rostral plate to

the apices of the submedian teeth of the telson;

CL refers to carapace length, measured on the midline, exclusive of rostral plate. Terms and measureRaymond B. Manning, Department of Invertebrate Zoology,

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

ments have been explained in earlier papers (see

Manning, 1969a).

The descriptive accounts vary in complexity depending on the taxon involved; thus a diagnosis

is given for a new Alima and a new Cloridopsis,

genera in which relatively few distinguishing characteristics are known. Often, characters mentioned

in generic diagnoses are not repeated in descriptions of species in that genus.

The following abbreviations are used to designate

repositories:

BMN'H British Museum (Natural History), London

MXHXP Museum National d'Histoire Naturelle, Paris

MRAC Musee Royal de 1'Afrique Centrale, Tervuren,

Belgium

MZC University Museum of Zoology, Cambridge, England

SME Station Marine d'Endoume, Marseille

SMF Natur-Museum und Forschungsinstitut Seiickenberg, Frankfurt am Main

USNM National Museum of Natural History, Smithsonian Institution, Washington (USNM = former

United States National Museum)

ZMB Zoologisches Museum an der Humboldt-Universita't zu Berlin

ZMH Zoologisches Institut und Zoologisches Museum,

Hamburg

ZSI Zoological Survey of India, Calcutta

ZSM Zoologisches Staatsammlung, Munich

ACKNOWLEDGMENTS.—A large part of this report

was made possible by B. Thomassin, M. PeyrotClausade, and R. Plante, Station Marine d'Endoume, Marseille, all of whom sent samples of

unidentified stomatopods, largely from Madagascar,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

for identification. H. Dingle, Department of Zoology, University of Iowa, and R. Caldwell, Department of Zoology, University of California, Berkeley, provided material from their field studies of

stomatopods in Thailand and freely discussed their

field observations. The following individuals also

provided working space and/or material for study

on loan at various times: P.L.G. Benoit, Musee

Royal de l'Afrique Centrale, Tervuren (directly

and indirectly through L. B. Holthuis, Rijksmuseum van Natuurlijke Historic Leiden); H.

Fechter, Zoologisches Staatsammlung, Munich; J.

Forest, Museum National d'Histoire Naturelle,

Paris; C. B. Goodhart, University Museum of Zoology, Cambridge; H.-E. Gruner and G. Hartwich,

Zoologisches Museum an der Humboldt-Universitat

zu Berlin; G. Hartmann, Zoologisches Institut und

Zoologisches Museum, Hamburg; R. W. Ingle,

Department of Zoology, British Museum (Natural

History), London; K. K. Tiwari and H. C. Ghosh,

Zoological Survey of India, Calcutta; and M. Tiirkay, Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt am Main.

Part of this study, including travel to Europe

in 1971 and India in 1972, was supported by the

Smithsonian Institution through its Research

Awards and Foreign Currency Programs.

I thank Anne Cohen and Horton H. Hobbs, Jr.,

for their careful review of the manuscript.

All of the illustrations were prepared by my

wife Lilly.

Family GONODACTYLIDAE Giesbrecht, 1910

Genus Gonodactylus Berthold, 1827

REMARKS.—Gonodactylus, one of the largest and

most complex genera in the Stomatopoda, comprises four distinct species groups, three of which

are restricted to the Indo-West-Pacific region. There

the genus is represented by the chiragra group,

composed of Gonodactylus chiragra (Fabricius,

1781) and its allies, including large species (TL

90 mm or more as adults) with large ocular scales

and three mid-dorsal keels or carinae on the telson

(note that G. viridis Serene, 1954, redescribed below, may belong here in spite of its small size and

ocular scales); the falcatus group, including G. falcatus (Forskal, 1775) and similar species, comprising small to large species (TL 30-60 mm or more

as adults) with small ocular scales and five middorsal keels on the telson; and the demanii group,

comprising G. demanii Henderson, 1893 and related species, encompassing small species (TL usually less than 50 mm as adults) with small ocular

scales and the telson variously ornamented with

longitudinal keels (usually three) and in some cases

with spinules. In some species of the latter group

the setation and shape of the uropod segments

may be highly modified. It is possible that this

group actually comprises species which should be

assigned to two groups, one with G. demanii, and

similar species, in which the telson and uropods

are usually modified, and one with G. affinis and

related species, in which the telson morphology

more closely resembles that of members of the

chiragra group.

The fourth aggregate of species in the genus,

the ocrstcdii group, including G. oersledii Hansen,

1895 and other American species, comprises small

to large species with small ocular scales, three middorsal keels on the telson, and an accessory or supplementary carina on the mesial side of the intermediate tooth of the telson.

Two species of the demanii group, G. affinis de

Man, 1902, and G. viridis Serene, 1954, are treated

below.

Gonodactylus affinis de Man, 1902

FIGURE 1

Gonodactylus chiragra var. affinis de Man, 1902:912.

Gonodactylm chiragra var. confinis de Man, 1902:912, pi. 27:

fig. 66.

Gonodactylus chiragra var. segregatus a Lanchester, 1903:448,

pi. 23: fig. 6.

Gonodactylus chiragra var. segregatus b Lanchester, 1903:448,

pi. 23: figs. 7, 7a.

Gonodactylus chiragra var. H. affinis.—Borradaile, 1907:211

[key], 212.

Gonodactylus chiragra var. K. confinis.—Borradaile, 1907:211

[key].

Gonodactylus segregatus.—Manning, 1968a:!>l, fig. 16.—Manning and Serene, 1968:114 [listed].—Manning, 1971:80 [key].

—Moosa, 1973:4 [listed], 8.

Gonodactylus confinis.—Manning, 1971:80 [key].

MATKRIAL.—Ternate, Molucca Islands, Indonesia; Kiikenthal, leg.; syntypes of Gonodactylus chiragra var. affinis de

Man: 1 $ . TL 21 mm; 2 9.TL 32-34 mm (SMF 5766). Same;

Kiikenthal, leg.; 1894; holotype of Gonodactylus chiragra var.

confinis de Man: 1 $ , TL 22 mm (SMF). Minikoi, Laccadive

Islands; reef; syntype of Gonodactylus chiragra var. segrega-
NUMBER 264

tus a Lanchester: 1 $, TL 18 mm (MZC). Goidu, Goifurfehendu Atoll, Maldive Islands; reef; syntypes of Gonodactylus

chiragra var. segregatus a Lanchester: 4 $, TL 10-15 mm

(MZC). South Male Atoll, Maldive Islands; 25 fms (46 m);

syntype of Gonodactylus chiragra var. segregatus a Lanchester: 15 , TL 11 mm (MZC). Hulule, Male Atoll, Maldive

Islands; reef; syntype of Gonodactylus chiragra var. segregatus a Lanchester: 1 juvenile, TL 8 mm (MZC). South

Nilandu Atoll, Maldive Islands; 25 fms (46 m); syntype of

Gonodactylus chiragra var. segregatus a Lanchester: 1 9 , TL

13 mm (MZC). South Nilandu Atoll, Maldive Islands; 19

fms (33 m); syntypes of Gonodactylus chiragra var. segregatus

b Lanchester: 2,$, TL 13-20 mm; 2$ , TL 14-15 mm; 1

juvenile, TL 9 mm (MZC).

One lot containing a single female from North Male Atoll

in 27-35 fms (49-67 m) could not be located in the collection at Cambridge.

9

FIGURE. 1.—Gonodactylus affinis de Man, 1902: a, female syntype, TL 34 mm, Ternate, sixth

abdominal somite, telson, and uropod. Gonodactylus confinis de Man, 1902, male holotype,

TL 22 mm, Ternate: b, rostral plate and ocular scales; c, sixth, seventh and eighth thoracic

somites, lateral view; d, sixth abdominal somite, telson, and uropod; e, submedian teeth of

telson, ventral view; /, uropod, ventral view. Gonodactylus segregatus Lanchester, 1903, female

syntype, TL 18 mm, Minikoi: g, sixth abdominal somite, telson, and uropod. (Setae omitted.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—The identity of the taxa described

as varieties affinis, confinis, and segregatus of Gonodactylus chiragra could not have been determined without examination of all of the types

involved. Illustrations of some of the types (Figure

1) are provided here. In G. affinis the accessory

median carinae of the telson are quite long and

often fuse posteriorly with the median carina. The

apices of the median, accessory median, and anterior submedian carinae each may be armed with a

small spinule. de Man (1902) described these spines

for G. affinis (a species that he believed to be distinct from G. confinis) in which the accessory median carinae are fused posteriorly with the highly

inflated median carina. His G. affinis was based on

four specimens, only three of which could be located, a male and two females. (The telson of one

of the females is shown herein: Figure la) with

relatively little inflation of the carinae of the telson, whereas his confinis was based on a male in

which the carinae are strongly inflated (Figure

Ib-f). Apparently both lots came from the same

collection made at Ternate, Molucca Islands. The

taxon described by Lanchester (1903) as G. chiragra

var. segregatus as from several localities in the western Indian Ocean is based on young specimens

and females with minimal inflation of the carinae

of the telson (Figure Ig).

The configuration of the mid-dorsal carinae on

the telson in this species superficially resembles

that found in G. falcatus (Forskal) and its allies.

Both de Man and Lanchester considered G. affinis

to be a transitional form between "typical" G. chiragra (Fabricius) and "typical" G. falcatus. It is,

in fact, not closely related to either but a member

of a group of small species that occur in the IndoWest-Pacific region.

This is a very small species; the largest individual available does not exceed 34 mm in total

length. Males as small as 22 mm may have welldeveloped secondary sexual characteristics.

DISTRIBUTION.—This species is known from Indonesia, where it was recorded from Ternate by

de Man (1902) and from several localities in the

Maluku region of Indonesia in 14-32 fms (26-59

m) (Moosa, 1973); Macclesfield Bank, South China

Sea (Lanchester, 1903); from Goidu, Goifurfehendu

Atoll, South Male Atoll in 25 fms (46 m), Hulule,

Male Atoll, North Male Atoll in 27-35 fms (49-64

m), South Nilandu Atoll in 19-25 fms (33-46 m),

and Minikoi, Laccadive Islands (all Lanchester,

1903); Seychelles, Amirante Island, Saya de Malha,

and Wasin, British East Africa (Wasini, Kenya)

(all Borradaile, 1907); and from Madagascar in

14 m and the Comoro Islands, intertidal zone

(Manning, 1968a).

Gonodactylus viridis Serene, 1954

FIGURES 2a-c

Gonodactylus chiragra var. viridis Serene, 1954:6, 7, 10, 74,

75, 76, fig. 13-3.

fGonodactylus chiragra.—Liu, 1975:188, figs. 3-4, 3-5 [part]

[not G. chiragra (Fabricius, 1781)].

MATERIAL.—KoPhuket Island, Thailand, Andaman Sea; in

shale and coral, shore to 1.5 m; R. Caldwell, leg.; Aug 1973:

3 $, TL 23-41 mm; 2 9. TL 33-43 mm (USNM 150597).

DESCRIPTION.—Rostral plate (Figure 2a) with

width slightly greater than length or length and

width subequal, apical spine relatively short; anterolateral angles broadly rounded, acute. Ocular

scales (Figure 2a) small, breadth of both less than

i/3 plate width, triangular, apices angled or rounded.

Sixth abdominal somite (Figure 26) with 6 carinae,

narrow in juveniles, inflated in adults, each usually

with apical tubercle or spinule. Telson (Figure 26)

appearing broad, length and width subequal or

width slightly greater, with 3 pairs of blunt marginal teeth, submedians with movable apices. Submedian, intermediate, and marginal carinae slender,

unarmed dorsally. Median and anterior submedian

carinae slender in juveniles and smaller females,

very inflated in large males (Figure 2c), unarmed

dorsally, usually unarmed posteriorly, each with

posterior tubercle in smaller specimens. Accessory

median carinae very short, not extending anteriorly beyond posterior third of median carina,

separate in juveniles, fusing posteriorly to form

anchor in adults. Knob indistinct or absent. Row of

submedian and 2 intermediate denticles present,

lateral denticles absent. Uropod (Figure 26) with

normal setation, proximal segment of exopod with

9-10 short, movable spines. Color completely

faded in available material.

MEASUREMENTS.—Males, TL 23-41 mm; females,

TL 33-43 mm. Serene (1954) had more than 300

specimens available, all less than 40 mm long, including females 35 mm long with eggs. Other
NUMBER 264

d

FIGURE 2.—Gonodactylus viridis Serene, 1954, female, TL 33 mm, Thailand: a, rostral plate and ocular scales; b, sixth abdominal somite, telson, and uropod. Male,

TL 41 mm, Thailand: c, sixth abdominal somite and telson. Gonodactylus chiragra

(Fabricius, 1781), female, TL 64 mm, Celebes: d, rostral plate and ocular scales.

(Setae omitted.)

measurements of male, TL 41 mm: carapace length

11.0; rostral plate length 2.8, width 3.0; fifth abdominal somite width 8.5; telson length 7.2, width

7.2.

REMARKS.—R. Serene named this species as a

variety of G. chiragra (Fabricius) in his account of

the biology of Vietnamese stomatopods published

in 1954, noting that a more complete account was

in preparation. It was named, figured, and distinguished from G. chiragra, which it resembles in

basic facies. Evidently it has not been recorded as

a distinct taxon in more recent taxonomic papers.

Apparently this species is easily recognized in the

field (R. Caldwell and H. Dingle, pers. comm.), but,

until the importance of the size of the ocular scales

was recognized, it was extremely difficult to separate it from juveniles of G. chiragra on the basis

of preserved material. It seems likely that some of

the records of G. chiragra from the Indo-Malayan

area should be referable to this species or to other

small species of the demanii group, such as G.

incipiens Lanchester, 1903 or G. micronesica Manning, 1971, which also have an unarmed telson

and normal setation on the uropods.

The G. chiragra-like telson and the small ocular

scales of G. viridis will distinguish it from all other

members of the demanii group. The small ocular

scales serve to distinguish it from G. chiragra even

in juveniles. The differences in the rostral plates

and ocular scales of this species and that of G.

chiragra (and the difference in ocular scale size

between members of the chiragra and demanii

groups) are shown in Figures 2a and 2d.

Gonodactylus viridis also differs from G. chiragra

in size: representatives of G. viridis are adult at a

length of 35 mm, and no specimens are known to

exceed 50 mm. Gonodactylus chiragra attains a

length of 105 mm (Kemp, 1913), and specimens

with total lengths of 70-90 mm are common.

It is with some hesitation that I assign the species to the demanii group, for the basic facies and

the telson of G. viridis and G. chiragra are very
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

similar. Obviously additional study of this species,

based on much more material than is now available,

is needed.

Liu (1975) may have included G. viridis with

material identified as G. chiragra in his account

of some stomatopods from the Xisha Islands, China.

His Figure 3 includes illustrations of the front and

telson of G. chiragra (Figures 3—1, 3-2) as well as

what appear to be those parts of G. viridis (Figures

3—4, 3-5). Unfortunately his account is in Chinese

and I am unable to clarify his records.

Serene found this species in blocks of coral in

the Bay of Nhatrang, and R. Caldwell collected

material in shale and coral in shallow water in

Thailand; it occurred together with G. chiragra

there.

DISTRIBUTION.—Known with certainty from the

Bay of Nhatrang, Viet Nam, and from KoPhuket

Island, Thailand, in the Andaman Sea.

Hoplosquilloid.es, new genus

DEFINITION.—Size small, TL not exceeding 25

mm. Rostral plate (Figure 3a) sharply trispinous.

Cornea (Figure 3a) elongate, not expanded laterally. Anterior margins of lateral plates of carapace

(Figure 3a) convex, extending anteriorly beyond

base of rostral plate. Ischiomeral articulation of

claw subterminal. Propodus of claw with proximal

movable spine. Dactylus of claw unarmed. Articulation of propodus and dactylus inflated. Mandibular palp present. Articulated anterolateral plates

of abdomen (Figure 36) present. Posterior margin

of sixth abdominal somite (Figure 3c) concave

medially. Telson (Figure 3c,d) of unusual shape,

ornamentation bizarre, dorsal and marginal armature arranged in circle around almost vertical

plane, anus opening in that plane above bases of

marginal teeth. Posteroventral margin of telson

produced into 2 primary pairs of long slender teeth,

submedians with movable apices, inner and outer

margins lined with long, slender, simple setae. Submedian denticles absent, replaced by simple setae.

Proximal segment of uropodal exopod (Figure

3c,/) extending beyond articulation with distal segment. Movable spines on outer margin of proximal

segment not markedly enlarged or strongly recurved. Endopod slightly curved mesially. Distal

segment of uropodal exopod and uropodal endopod (Figure 3c,/) with strong fixed spines on mesial

margin.

TYPE-SPECIES.—Hoplosquilloides coronatus, new

species.

ETYMOLOGY.—The name is derived from the

Greek -oides, meaning "like," in combination with

the generic name Hoplosquilla. The gender is

masculine.

REMARKS.—Hoplosquilloides is unique among

stomatopods in having the anus situated dorsally

(Figure 3c) rather than ventrally on the telson.

This may well be an adaptation for burrowing,

but this remains speculative, for live animals have

not yet been observed. Hoplosquilloides shows

closest affinities with Hoplosquilla Holthuis, 1964,

represented by a single species from Ceylon, in

that the uropod is armed with sharp, fixed spines.

Hoplosquilloides differs from Hoplosquilla in that

the mandibular palp is present and in the shape

and armature of the telson, with the anus situated

dorsally rather than ventrally. The telson in Hoplosquilla is typically gonodactylid, not highly modified as in this new genus.

Hoplosquilloides coronatus, new species

FIGURE 3

MATERIAL.—St. Gilles Reef, Reunion Island; outer slope,

35 m; M. Clausade, leg., sta RE-74-22; 1974; paratype: 1 9 ,

TL 12 mm (SME). St. Gilles Reef, Reunion Island; outer

slope, 35 m; M. Clausade, leg., sta RE-74-25; 1974; holotype:

19, TL 18 mm (MNHNP). Same; reef flat; M. Clausade,

leg., sta RE-74-55; 1974; paratype: 1 $ , TL 18 mm (USNM

156240). Grand Recif, Tulear, Madagascar; external slope,

21 m; B. Thomassin, leg., sta 230; 3 Oct 1969; paratype: 1 $ ,

TL 13 mm (MNHNP). Grand Recif, Tulear, Madagascar;

Antseteky transect; 21-33 m; Pichon, leg., sta D-8; 2 Sep

1971; paratype: 1 $, TL 23 mm (SME). Grand Recif, Tulear, Madagascar; Antseteky transect, 31 m, hard bottom;

Pichon leg., sta. D-13; 2 Sep 1971; paratypes: 1 $, TL 17

mm; 2 $ . TL 17-225 mm (USNM 156168).

DESCRIPTION.—Ocular scales (Figure 3a) low but

distinct, rounded or flattened dorsally, separate

mesially. Cornea (Figure 3a) elongate, diameter

greater than that of stalk.

Rostral plate (Figure 3a) sharply trispinous,

median spine noticeably longer than anterolaterals.

Mandibular palp three-segmented. 5 epipods

present.
NUMBER 264

FIGURE 3.—Hoplosquilloides coronatus, new genus, new species, female paratype, TL 22.5 mm,

Grand Recif: a, anterior portion of body; b, first abdominal somite, lateral view; c, sixth

abdominal somite and telson; d sixth abdominal somite and telson, lateral view; e, uropod,

dorsal view; /, uropod, ventral view. (Setae omitted in e, /.)

Exposed thoracic and anterior 5 abdominal somites smooth, unarmed, lacking sharp dorsal carinae, abdominal somites each with marginal carina.

Sixth abdominal somite (Figure 3c) with 6 longitudinal bosses, submedians each with posterior

spinule, intermediates bluntly pointed, laterals

produced into slender posterolateral spine. Anterior surface of lateral bosses irregular, appearing

eroded.

Telson (Figure 5c,d) much broader than long

with short basal portion connected to long marginal teeth by curved vertical surface, appearing

semicircular or oval in posterior view, margin

framed by short, rounded bosses dorsally and laterally, by long marginal teeth ventrally. Each of dorsal bosses with at least 1 posterior tubercle or spine,

some bosses apically bifurcate or with secondary

spinules around apical spine. Short but distinct

bluntly pointed lateral tooth present, carinate laterally. Posterior margin with ventral 2 pairs of

projections longest, submedians with movable

apices, intermediates slightly shorter, with fixed

apices. Submedian, intermediate, and lateral denticles absent. Inner and outer margins of submedian
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and intermediate marginal teeth completely lined

with single series of long, simple setae. Anus (Figure 3c) visible in dorsal view, situated posterodorsally rather than ventrally, encircled dorsally and

laterally by ridge connecting inner margins of

submedian teeth.

Proximal segment of uropod (Figure 3e,f) with

sharp, curved carina dorsally, terminating in sharp

spine, 1-2 tubercles present dorsally lateral to carina. Proximal segment of uropodal exopod with

11-12 movable spines laterally, distalmost slightly

recurved but not enlarged, and with fixed spine

ventrally (Figure 3/) overhanging articulation with

distal segment. Inner margin of proximal segment

of exopod (Figure 3e) smooth, lacking setae, with

sharp spine at inner, distal angle. Distal segment

of uropodal exopod (Figure $et,f) with normal setation laterally, lacking setae but with 3-5 fixed

spines mesially, and dorsally with fixed spines on

short carina. Uropodal endopod (Figure 3e) with

normal setation (margin evenly fringed) on outer

margin, inner margin lacking setae but with strong

proximal spine and several slender fixed spines

distally, margin smooth between spines.

COLOR.—Largely faded in preservative, but with

tip of rostral plate red, anterior appendages mottled with dark pigment, and eyestalks brown with

dark mottling. Carapace with row of dark spots

across posterior third. Dactylus of claw orange or

pink, less orange pigment on propodus and at

merocarpal articulation. Posterior 3 thoracic somites each with dark spots arranged in band across

somite, most prominent on sixth. All 6 abdominal

somites with pair of submedian black spots, first

somite also with larger, extra pair nearer to posterior margin, and first and fifth somites with median spot. Telson with anterior pair of small, dark

spots.

MEASUREMENTS.—Males, TL 13-17 mm; females,

TL 12-23 mm. Other measurements of female holotype, TL 18 mm: carapace length 3.8; fifth abdominal somite width 3.0; telson length 2.2, width 2.7.

REMARKS.—The specimens examined are remarkably uniform, the major variability observed being

on the ornamentation of the telson. In the smaller

specimens (TL 12-13 mm), the lateral projections

are much less ornate than in the larger examples.

There is also some minor variation in the size and

position of the fixed spines on the margins of the

uropod. The telson in all of the specimens appears

to have a basic pattern of a median (dorsal) and

seven lateral marginal projections, somewhat obscured by secondary ornamentation of the major

projections.

ETYMOLOGY.—The specific name is from the

Latin corona, "crown," and -atus, "having the

shape of," alluding to the shape of the telson.

DISTRIBUTION.—Known only from Reunion Island

and the Grand Recif, Tulear, Madagascar. It lives

on coral reefs, from the reef flat to a depth of 35

m on the slope.

Family LYSIOSQUILLIDAE Giesbrecht, 1910

Genus Heterosquilla Manning, 1963

Heterosquilla tricarinata (Claus, 1871)

FK;URF. 4

Coronis tricarinata Claus, 1871:128, 139 [pagination from

Holthuis, 1967a; pp. 21, 31 on separate].

Coronis spinosa Wood-Mason, 1875:232.

Heterosquilla (Heterosquilla) tricarinata.—Holthuis, 1967a: 11

[complete synonymy].—Manning and Serene, 1968:115

[listed].

Heterosquilla (Heterosquilla) spinosa.—Holthius, 1967a: 11

[complete synonymy].—Manning and Serene, 1968:115

[listed].

MATERIAL.—Port Blair, Andaman Islands; J. Wood-Mason,

don.; lectotype of Coronis spinosa: 1 $, TL 68 mm (ZSI

3082/5).

REMARKS.—In 1966 I suggested that H. spinosa

from the Andaman Islands probably was distinct

from H. tricarinata from New Zealand. This was

based on differences between the accounts of material from the two areas, the most important being

the absence of the inner spine on the basal prolongation of the uropod in H. spinosa. Examination of the lectotype of that species in 1972 revealed

that the inner spine was broken on one side (Figure 4e), and for this reason it was shown as an

inconspicuous lobe by Kemp (1913, pi. 8: fig. 94).

I can find no other differences between the specimen from the Andaman Islands and accounts of

material from New Zealand, and I believe that

Coronis spinosa Wood-Mason, 1875 must be considered a synonym of Coronis tricarinata Claus,

1871. The specimen figured here is the lectotype

of Coronis spinosa.
NUMBER 264

FIGURE 4.—Heterosquilla tricarinata (Claus, 1871), female lectotype, TL 68 mm, Andaman

Islands: a, base of third walking leg, oblique ventral view; b, telson, dorsal view; c, posterior

margin of telson, ventral view; d, basal part of right uropod, ventral view (basal prolongation

broken); e, basal part of left uropod, ventral view (inner spine of basal prolongation broken).

Keppelius, new genus

DEFINITION.—Size very small, maximum length

less than 30 mm. Body smooth, strongly depressed,

loosely-articulated. Cornea (Figure 5a) subglobular, expanded laterally beyond stalk and set slightly

obliquely on it. Rostral plate (Figure 5a) subrectangular, anterior margin with three projections.

Antennal protopod with at most 1 ventral papilla.

Carapace short, narrowed anteriorly, carinae and

spines absent, cervical groove scarcely indicated on

lateral plates. Exposed thoracic somites lacking

carinae; eighth thoracic somite lacking median

ventral keel. Mandibular palp absent, 5 epipods

present. Claw (Figure bb) stout, dactylus with 7

or more teeth, outer margin with basal notch;

propodus stout, fully pectinate, with 4 movable

spines proximally; dorsal ridge of carpus undivided, terminating in acute lobe; merus slender,

unarmed; ischiomeral articulation terminal; ischium three-fourths length of merus, unarmed.

Endopods of pereopods two-segmented, distal segment subcircular on anterior 2 legs, more ovate

on third. Abdomen without longitudinal carinae

or spines, articulated anterolateral plates small;

sixth abdominal somite (Figure 5c) smooth dorsally, with posterolateral spines and curved, ventrally-directed process anterior to each uropod. Telson (Figure bc,d) broader than long, posterior part

of surface covered with posteriorly-directed spines,

marginal armature, obscured by dorsal spines, including long, movable submedian teeth and other

fixed teeth and denticles. Basal segment of uropod

(Figure 5c) with sharp inner and outer edges, inner

terminating in distal, dorsal spine; proximal segment of uropodal exopod (Figure 5r) with spatulate spines on outer margin, rounded lobe with

stiff setae on inner, distal margin; endopod (Figure

5c) oval, with strong proximal fold on outer margin; inner spine of basal prolongation of uropod

the longer (Figure be).

TYPE-SPECIES.—Lysiosquilla hystricotelson Barnard, 1958.

ETYMOLOGY.—It is a pleasure to name this genus

for the late Keppel H. Barnard, South African

Museum, Capetown. It is based on the only stomatopod described by him. The gender is masculine.

REMARKS.—Keppelius shares numerous features

with Acanthosquilla Manning, 1963, Nannosquilla
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Manning, 1963, and Pullosquilla, new genus. It

differs from Acanthosquilla in the configuration of

the dorsal spines on the telson. In Acanthosquilla

these spines are arranged in a transverse row across

the telson surface, whereas in Keppelius there are

many more spines covering a vertical shelf just

above the posterior margin of the telson. Keppelius

lacks distinguishable primary marginal teeth (intermediate, lateral, and marginal) on the telson,

for all of the fixed marginal teeth and denticles

are subequal in size. Finally, in Keppelius the movable outer spines on the uropodal exopod are

spatulate rather than slender. Keppelius differs

from Pullosquilla in lacking an ischial spine on

the claw and in having a rectangular rather than

a triangular rostral plate. Keppelius agrees with

Nannosquilla in basic facies, but lacks an ornamental eave above the marginal armature of the

telson and also differs in having five rather than

four epipods, a dorsal spine on the antennal protopod, and one antennal papilla.

Keppelius contains only the type-species; it is

redescribed below.

Keppelius hystricotelson (Barnard, 1958)

FIGURE 5

Lysiosquilla hystricotelson Barnard, 1958:20, fig. 7.—Manning,

1963:319 [listed; transferred to Nannosquilla].

Nannosquilla hystricotelson.—Holthuis, 1967a:23 [synonymy];

1967b:25, 40 [listed].—Tirmizi and Manning, 1968:5 [listed].

—Manning and Serene, 1968:116 [listed].—Manning, 1969b:3

[list; key].—Makarov, 1971:154 [listed].—Thomassin, 1974:

303.

MATERIAL.—Mombasa, Kenya; on reef exposed at S entrance

to harbor; poisoned; A. J. Bruce, leg; Anton Bruun Cr. 9;

15 Nov 1964: 1 $, TL 23 mm (USNM 168779). Tulear,

Madagascar; B. Thomassin, leg., sta 469: 1 $, CL 3.8 mm

(MNHNP). Near Songoritelo Reef, Madagascar; B. Thomassin, leg., sta 444; 19 Oct 1962: 1 & , TL 19 mm; 1 9 , TL 20

mm (USNM 139073). Nosy Be, Madagascar; E beach; infra

littoral, in grass; B. Thomassin, leg., sta 851; 13 Jul 1972:

1 $, TL 20.5 mm; 19 TL 23 mm (USNM 156175); and 1 $,

TL 18 mm; 19 TL 24 mm (SME). Nosy Be, Madagascar;

W beach; in grass; B. Thomassin, leg., sta 880; 14 Jul 1972:

19,1 carapace (MNHNP).

DIAGNOSIS.—Ocular scales fused into bilobed

plate. Antennal protopod with sharp antero-internal spine (Figure 5a). Antennular peduncle less

than half as long as carapace. Antennal scale (Figure 5a) about 14 as long as carapace. Dactylus

of claw (Figure 56) with 7-10 (usually 8) teeth,

proximal notch on outer margin flanked by sharp

proximal and obtuse distal prominences. Basal segments of pereopods unarmed. Sixth abdominal

somite (Figure bd) with 2 large, posteriorly directed

spines on ventral surface; marginal armature of

telson (Figure bc,d), on each side of midline, consisting of: 5-9 (usually 6-7) submedian denticles

(arranged in inverted, V-shaped row), 1 movable

submedian tooth, and 7 fixed lateral teeth and

denticles. Proximal segment of uropodal exopod

(Figure bc,e) with 3^1 blunt, movable spines on

outer margin, distal 2 spatulate, and 1 stiff, spinelike seta on inner, distal angle.

COLOR.—Faded in most specimens examined, but

some with traces of light anterior and darker posterior bands on carapace and with small chromatophores scattered over remainder of body in no

particular pattern.

MEASUREMENTS.—Males, TL 18-20.5 mm; females, TL 20-24 mm. Barnard's type, a male, was

25 mm long. Other measurements of a female, TL

23 mm: carapace length 4.0; cornea width 0.7; rostral plate length 1.5, width 1.6; antennal scale

length, 0.9; fifth abdominal somite width 4.0; telson length 2.2, width 3.4.

REMARKS.—This species, with its broad rostral

plate, produced anteriorly into three sharp projections, and characteristically ornamented telson, can

readily be distinguished from all other lysiosquillids. It may be confused with species of Acanthosquilla that have more than 5 spines on the dorsal surface of the telson, but in them the telson

bears fewer dorsal spines as well as a different

configuration of marginal teeth. The ventral antennal papilla is not visible in all specimens.

Thomassin (1974) found this species in sea grass

beds of the fringing reef flat at Songoritelo, Madagascar. Like the species of Pullosquilla discussed

below, K. hystricotelson is a shallow water species

living on reef flats in grass beds on sand. The specimen reported by Holthuis (1967b) came from dead

coral near Massawa, Ethiopia.

"DISTRIBUTION.—Western Indian Ocean, where it

has been recorded from a few localities between

the type-locality, Delagoa Bay, Mozambique (Barnard, 1958) and Massawa, Ethiopia (Holthuis,

1967b), including Songoritelo, Madagascar (Thomassin, 1974), and the other localities reported

above: Mombasa, Kenya, and Tulear and Nosy Be,

Madagascar.
NUMBER 264 11

FIGURE 5.—Keppelius hystricotelson (Barnard, 1958), female, TL 23 mm, Mombasa: a, anterior

part of body; b, carpus, propodus, and dactylus of claw; c, sixth abdominal somite, telson,

and uropod, dorsal view; d, sixth abdominal somite and telson, ventral view; e, uropod,

ventral view. (Setae omitted.)

Genus Neocoronida Manning, 1976

Neocoronida was established by Manning (1976a)

for two species of lysiosquillids that previously had

been assigned to Coronida Brooks, 1886: N. trachurus (von Martens, 1881), from the Palau Islands

and Mauritius in the Indo-West-Pacific region, and

N. cocosiana (Manning, 1972), from Cocos Island

in the eastern Pacific. Examination of two syntypes

of N. trachurus has revealed that they are not

conspecific. Neocoronida trachurus sensu stricto,

from the western Indian Ocean, and N. martensi,

new species, from the Palau Islands, are described

and illustrated below.

Key to Species of Neocoronida

1. Dorsal tubercles of telson all sharp, stellate N. trachurus (von Martens, 1881)

Dorsal tubercles of telson all rounded dorsally 2

2. Rostral plate pentagonal, with slight apical projection. Ventral surface of telson tuberculate

2V. martensi, new species

Rostral plate ovate, rounded anteriorly. Ventral surface of telson smooth

N. cocosiana (Manning, 1972)
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Neocoronida martensi, new species

FIGURE 6

Gonodactylus trachurus von Martens, 1881:93 [part].

MATERIAL.—Palau Islands, Caroline Islands, western Pacific;

Piitze, leg., holotype [paralectotype of Neocoronida trachurus

(von Martens, 1881)]: 1 9, TL 45 mm (ZMB 6159).

DESCRIPTION.—Size small, only known specimen

45 mm long.

Eye (Figure 6a) elongate, triangular, cornea

noticeably broader than and set obliquely on stalk.

Eye extending about to end of first segment of

antennular peduncle. Ocular scales (Figure 6a) low,

broad, separate. Anterior margin of ophthalmic

somite produced into median spinule.

Antennular peduncle about % as long as carapace. Antennular processes (Figure 6a) produced

into blunt projections, appearing sharp in dorsal

view, visible lateral to rostral plate. Shorter ramus

of ventral flagellum with 21 segments in holotype.

Antennal scale (Figure 6a) more than half as

long as carapace. Proximal segment of antennal

peduncle longer than distal segment, extending

beyond eye. Protopod with blunt anterointernal

projection and 1 ventral papilla.

Rostral plate (Figure 6a) pentagonal, broader

than long, unarmed. Lateral margins slightly convex, anterolateral angles broadly rounded, concave

anterior margins converging on obtusely-pointed

apex. Low obscure median carina present.

Dactylus of claw with 4 teeth, outer margin

inflated, notched basally. Dorsal ridge of carpus

b

FIGURE 6.—Neocoronida martensi, new species, female holotype, TL 45 mm, Palau Islands:

a, anterior part of body; b, posterior part of body; c, tubercle of telson, lateral view; d, telson,

ventral view; e, uropod, ventral view. (Setae omitted.)
NUMBER 264

13

low, terminating in blunt projection proximal to

distal margin.

Mandibular palp present. 5 epipods present.

Lateral process of fifth thoracic somite rounded,

compressed anteroposteriorly, appearing acute in

dorsal view. Fifth somite also with small ventral

spinule on each side. Lateral processes of sixth and

seventh thoracic somites broadly rounded. Eighth

thoracic somite without noticeable median ventral

keel. Endopods of perepods slender, two-segmented.

Anterior 4 abdominal somites smooth dorsally,

lacking tubercles or spines. Anterior portion of fifth

somite smooth, remainder of surface (Figure 6b)

with rounded dorsal tubercles, more numerous

laterally. Fifth somite with 4 pairs of longitudinal

grooves, lateralmost longest, between margin and

longitudinal carina above margin; posterior margin

lined with subequal tubercles or spinules, posterolateral angles lacking spines. Sixth somite (Figure

6b) completely covered with stalked, plate-like

tubercles, margins irregular, surface smooth (Figure

6c); lateral margin forming broad, carinate ridge,

and with blunt posterolateral projection; posterior

margin lined with spinules; acute but blunt projections present ventrolaterally anterior to articulation of each uropod.

Telson (Figure 6b,d) broader than long, posterior

margin strongly convex, dorsal surface completely

covered with stalked, plate-like tubercles (Figure

6c), margins irregular, surface smooth. Posterior

armature of telson as illustrated. Ventral surface of

telson (Figure 6d) with median patch of tubercles

between anal pore and posterior margin.

Uropod (Figure 6b,e) broad, flattened, anterior

margin of basal segment carinate. Dorsal surface of

basal segment ornamented with numerous rounded

tubercles, all simple, and with short dorsal carina

bearing minute apical spine. Each segment of uropod with dorsal tubercles, tubercles also present on

ventral surface of distal segment of exopod. Outer

margin of proximal segment of exopod with 9

short, movable spines, posteromesial lobe of proximal segment not markedly projecting, lacking stiff

setae. Distal segment of uropodal exopod longer

than proximal. Endopod curved mesially, lacking

strong proximal fold laterally. Spines of basal prolongation (Figure 6e) flattened, inner longer, with

3 slender, fixed spines on inner margin.

COLOR.—Completely faded in holotype.

MEASUREMENTS.—Unique female holotype, TL

45 mm. Other measurements: carapace length 7.4;

cornea width 2.4; antennular peduncle length 6.1;

antennal scale length 4.1; rostral plate length 1.6,

width 2.1; fifth abdominal somite width 9.4; telson

length 5.6, width 9.1.

REMARKS.—This species is known only from the

holotype, which also is the paralectotype of Neocoronida trachurus (von Martens, 1881). Neocoronida

martensi resembles iV. trachurus in most respects,

differing in that the raised, plate-like tubercles on

the sixth abdominal somite and telson are completely smooth dorsally, and the dorsal tubercles

of the fifth abdominal somite and uropods are

larger, rounder, and smoother. The new species

more closely resembles N. cocosiana (Manning),

from Cocos Island in the Eastern Pacific, especially

in the form of the plate-like tubercles, but that

species has a rounded rostral plate and the ventral

surface of the telson is smooth.

ETYMOLOGY.—The species is named for the German carcinologist E. von Martens, who discovered

N. trachurus and considered the specimen herein

designated as the holotype of N. martensi as one of

its members.

DISTRIBUTION.—Known only from the typelocality, the Palau Islands. The depth range and

habitat have not been recorded.

Neocoronida trachurus (von Martens, 1881)

FIGURE 7

Gonodactylus trachurus von Martens, 1881:93 [part].

Coronida trachurus.—Holthuis, 1967a:8 [references to 1967];

1967b:40 [Red Sea references].—Makarov, 1971:153 [listed].

Coronida trachura.—Manning and Serene, 1968:115 [listed].—

Tirmizi and Manning, 1968:4 [listed].

MATERIAL.—Red Sea; Jousseaume, leg.; 1898: 1 $, TL 33

mm (MNHNP). Between Grande Pass and Johny Channel,

Aldabra Island; 42 m; 23 May 1953: 1 $, TL 36 mm (USNM

169780); 1 9 , TL 36 mm (MNHNP). Mauritius; G. Schneider, leg.; lectotype: 1 $, TL 47 mm (ZMB 6365). Mauritius; no other data: 2 $ (1 broken), TL 32 mm (MNHNP).

Mauritius; no other data: 4 specimens, not measured (ZSI

4271/4).

DESCRIPTION.—Size small to moderate, total length

less than 50 mm. Eye (Figure la) elongate, triangular, cornea bilobed, noticeably broader than and

set obliquely on stalk. Eyes extending about to end

of first segment of antennular peduncle. Ocular

scales low, broad, separate. Anterior margin of
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Neocoronida trachurus (von Martens, 1881), male lectotype, TL 47 mm, Mauritius;

a, anterior part of body; b, carpus, proporius, and dactylus of claw; c, lateral processes of

fifth, sixth, and seventh thoracic somites; d, posterior part of body; e, tubercle of telson,

lateral view; /, telson, ventral view; g, uropod, ventral view. (Setae omitted.)

ophthalmic somite produced into median spinule

or tubercle.

Antennular peduncle about 2/3 as long as carapace. Antennular processes (Figure 7a) produced

into blunt projections, appearing sharp in dorsal

view, visible lateral to rostral plate. Shorter ramus

of antennular fiagellum with 16-19 segments.

Antennal scale about half as long as carapace.

Proximal segment of antennal peduncle longer

than distal, extending beyond eye. Antennal protopod with blunt antero-internal projection and 1

ventral papilla.

Rostral plate (Figure 7a) pentagonal, broader

than long, unarmed. Lateral margins straight or
NUMBER 264 15

slightly concave, anterolateral angles broadly

rounded, concave anterior margins converging on

obtusely-pointed apex. Low median carina present

or absent.

Dactylus of claw (Figure 7b) with 4 teeth, outer

margin inflated, notched basally. Dorsal ridge of

carpus low, terminating in blunt projection proximal to distal margin.

Mandibular palp present. 5 epipods present.

Lateral process of fifth thoracic somite (Figure

7c) rounded, compressed anteroposteriorly, appearing acute in dorsal view, Fifth somite also with

small ventral spinule or rounded tubercle on each

side. Lateral processes of sixth and seventh somites

(Figure 7c) broadly rounded, that of sixth somite

with rounded tubercles mesial to lateral margin.

Eighth somite with low, rounded median prominence ventrally, lacking distinct ventral keel. Endopods of pereopods slender, two segmented.

Anterior 4 abdominal somites smooth dorsally,

lacking tubercles or spines. Anteromedian portion

of fifth somite (Figure 7d) smooth, remainder of

surface completely covered with variously-shaped

tubercles; several short, longitudinal carinae present above lateral margin, largest fusing posteriorly

into marginal carina; posterior margin (Figure Id)

lined with sharp tubercles or spinules, increasing

in size mesially. Sixth somite (Figure Id) completely

covered with stalked, stellate, or spinous tubercles

(Figure 7e); lateral margin forming irregular carina

terminating in blunt posterolateral projection;

posterior margin (Figure Id) lined with sharp

tubercles and spinules; blunt projections present

ventrolaterally anterior to articulation of each

uropod.

Telson (Figure 7d,f) broader than long, posterior

margin strongly convex, dorsal surface completely

covered with stalked, stellate tubercles (Figure 7e).

Posterior armature of telson as illustrated (Figure

7d,f). Ventral surface of telson (Figure 7/) with

median patch of tubercles between anal pore and

posterior margin.

Uropod (Figure 7d,g) broad, flattened, anterior

margin of basal segment carinate, dorsal surface

covered with simple and stellate tubercles and with

short dorsal carina terminating in blunt spine. Each

segment of uropod with dorsal tubercles, tubercles

also present on ventral surface of distal segment of

exopod. Outer margin of proximal segment of

exopod (Figure 7d) with 8-10 short, movable spines,

posteromesial lobe of proximal segment not markedly projecting, stiff setae not discernible. Distal

segment of uropodal exopod (Figure 7d) longer

than proximal. Endopod (Figure 7g) curved mesially, lacking strong proximal fold laterally. Spines

of basal prolongation (Figure 7g) flattened, inner

longer, with 1-4 slender, fixed spines on inner

margin.

COLOR.—Faded in most specimens, but in those

from Aldabra body completely covered with numerous small dark chromatophores; pigment concentrated on posterolateral angles of carapace,

median area of the sixth thoracic somite, and anterolateral plates of abdomen. Each somite with

faint median and 2 lateral dark patches, body

appearing striped in dorsal view.

MEASUREMENTS.—Males, TL 32-47 mm; only

female measured, TL 36 mm. Other measurements

of male lectotype, TL 47 mm: carapace length 9.0;

cornea width 2.5; antennular peduncle length ca.

6.6; rostral plate length 2.0, width 2.5; fifth abdominal somite width 10.7; telson length 6.5, width 10.6.

REMARKS.—Neocoronida trachurus, which apparently is restricted to localities in the western Indian

Ocean, is readily distinguishable from the other

two species of the genus by the presence of sharply

stellate stalked tubercles on the sixth abdominal

somite and telson; the other two species both have

stalked tubercles which are rounded dorsally.

Although von Martens (1881:93) mentioned two

specimens from Mauritius, only one, a female

47 mm long, could be located. It is herein selected

as the lectotype of Gonodactylus trachurus. The paralectotype of G. trachurus is also the holotype of

Neocoronida martensi, described above.

DISTRIBUTION.—Western Indian Ocean, where it

has been recorded from the Red Sea, Aldabra

Island, and Mauritius. Its depth range and habitat

have not been recorded.

Genus Parvisquilla Manning, 1973

Parvisquilla Manning, 1973:209.

DEFINITION.—Size very small, maximum length

less than 20 mm. Body smooth, compact, depressed.

Eye (Figure 8a) elongate, cornea faintly bilobed.

Rostral plate (Figure 8a) very short, broadly

rounded anteriorly. Antennular somite (Figure 8a)

elongate, not overreached by rostral plate. Antennal
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

protopod lacking discernible papillae. Carapace

strongly narrowed anteriorly, carinae and spines

absent, cervical groove not discernible, gastric

grooves very fine, poorly-marked. Exposed thoracic

somites (Figure Sd) lacking longitudinal carinae;

eighth thoracic somite lacking median ventral keel.

Mandibular palp absent, 2 epipods present. Propodi of third and fourth (Figure 86) thoracic appendages broader than long, lacking distinct ventral

beading or ribbing. Claw (Figure 8c) stout, dactylus

with 4-5 teeth, outer margin of dactylus inflated

basally, with 3 proximal projections; propodus stout,

proximal margin incompletely pectinate, with 3

movable spines proximally; dorsal ridge of carpus

undivided, terminating in sharp spine; merus stout,

unarmed; ischium much shorter than merus,

unarmed. Endopods of pereopods 1-segmented,

slender. Abdomen depressed but compact, smooth;

articulated anterolateral plates absent; sixth abdominal somite (Figure Se) ornamented with irregular ridges and tubercles. Telson (Figure 8^,/)

broader than long, completely covered with raised,

rounded tubercles or irregular ridges; marginal

armature of telson including submedian teeth with

movable apices and other fixed teeth and denticles.

Uropod (Figure Se,g) stout, proximal segment with

prominent outer ridge; outer margin of proximal

segment of exopod completely lined with blunt

spines, inner margin setose; endopod lacking prominent fold proximally on outer margin; basal prolongation (Figure 8g) flattened, produced into 2

short blunt spines, inner longer.

TYPE-SPECIES.—Squilla multituberculata Borradaile, 1898.

REMARKS.—This genus was established by me

for two species previously assigned to Coronida,

and, through an error in interpretation on my part,

was transferred from the family Lysiosquillidae to

the family Squillidae. The propodus of the third

and fourth maxillipeds, in situ, appeared to be

squillid in shape, slender and lacking the beading

or ribbing characteristic of lysiosquillids. Subsequently I dissected one of the maxillipeds of P.

multituberculata to prepare the illustration presented here (Figure 8b) to find that it was typically

lysiosquillid in shape but lacked the ventral ribbing typical of members of the Lysiosquillidae.

Further, a better light source revealed that the gastric grooves, although very faint, are indeed present.

Borradaile (1898, pi. 6: fig. 7), Edmondson (1921,

fig. 2) and Liu (1975, pi. 1: fig. 1) all omitted the

gastric grooves in their figures.

Parvisquilla, which appears to be related to

Coronida and Neocoronida, must be referred to the

Family Lysiosquillidae. It agrees with these genera

in having a stout claw with a basally-inflated dactylus. Its distinctive features include the greatly

elongated antennular somite, the very short rostral

plate, the exterior projections on the dactylus of

the claw, and the suppressed mandibular palp.

Two species are assigned to Parvisquilla, and one

of these, P. multituberculata, is redescribed below.

The other species, P. sinuosa (Edmondson, 1921),

is known only from Hawaii.

Key to Species of Parvisquilla

Telson covered with irregular pattern of linear, curved, and scroll-like carinae

P. sinuosa (Edmondson, 1921)

Telson covered with erect, rounded tubercles P. multituberculata (Borradaile, 1898)

Parvisquilla multituberculata (Borradaile, 1898)

FIGURE 8

Squilla multituberculata Borradaile, 1898:38, pi. 6: fig. 7;

1899:403.—Manning. 1963:322 [listed]; 1973:299 [listed].

Coronida multituberculata.—Kemp, 1913:132.—Gravier, 1927:

36 [discussion].—Holthuis, 1967a:8 [synonymy].—Manning

and Serene, 1968:115 [listed].

Squilla tubercitlata.—Gravier, 1927:36 [error for S. multituberculata}.

Parvisquilla multituberculata.—Manning, 1973:299 [discussion].

Parvisquilla xishaensis Liu, 1975:183, 196, pi. 1: figs. 1-6.

MATERIAL—Eva Island, Tonga Islands; 21°22'S, 174°56'W;

Albatross, leg.; 1899-1900: 1$ , TL 13 mm (USNM 125330).

Apia, Samoa; no other data: 2$ , TL 11-12 mm (ZSM).

Moorea, Society Islands; reef flat; M. Clausade, leg., sta

MO-73-15; 1973: 1 $, TL 13 mm (SME).

DESCRIPTION.—Size very small, TL of adults 13

mm or less. Body strongly depressed, surface smooth.

Eye (Figure 8a) small, cornea bilobed, set

obliquely on and not markedly broader than stalk.
NUMBER 264 17

FIGURE 8.—Parvisquilla multituberculata (Borradaile, 1898), female, TL 13 mm, Tonga Islands:

a, anterior part of body; b, propodus and dactylus of fourth maxilliped; c, claw; d, lateral

processes of fifth, sixth, and seventh thoracic somites; e, sixth abdominal somite, telson, and

uropod; /, margin of telson, ventral view; g, uropod, ventral view. (Setae omitted, except b.)

Eyes extending almost to end of first segment of

antennular peduncle. Ocular scales (Figure 8a)

broad, truncate, separate mesially. Anterior margin

of ophthalmic somite produced into median spinule.

Antennular peduncle as long as carapace. Antennular processes produced into broad, blunt lobes

directed almost anteriorly. Antennular somite

(Figure 8a) elongate, much longer than rostral

plate.

Antennal protopod elongate, extending beyond

antennular processes. Antennal scale very short,

ovate. Proximal segment of antennal peduncle

longer than distal.

Rostral plate (Figure 8a) triangular, about twice

as wide as long, very short, not extending to level

of articulation of antennal peduncle, lacking

carinae or spines.

Carapace very small, strongly narrowed anteriorly, lacking carinae and spines. Gastric grooves

present, scarcely visible at magnifications of less

than X 50. Anterior margins almost straight, anterolateral angles bluntly rectangular, not projecting

anteriorly. Carapace not concealing chelae, epipods,

or fifth thoracic somite.

Claw (Figure 8c) stout, large, almost all of ischium

and merus visible in dorsal view. Dactylus sinuous,

inflated basally, armed with 4 teeth; outer margin

with 3 angular projections proximally, proximal

largest, distalmost more obtuse than basal 2. Propodus deep, proximal s/4 pectinate. Dorsal ridge of

carpus terminating in sharp tooth.

Mandibular palp absent. 2 epipods present.

Exposed thoracic somites (Figure 8d) lacking

dorsal carinae or spines. Lateral process of fifth

somite forming a rounded lobe, oblique to body

line, compressed antero-posteriorly; blunt ventral
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spine also present on each side. Lateral processes

of sixth to eighth somites broad, truncate, flattened

or rounded laterally. Pereopods stout, short, endopods composed of 1 segment.

Abdomen smooth, broad, depressed, looselyarticulated, anterior 5 somites lacking spines or

longitudinal carinae. Anterolateral plates of abdomen absent. Fifth somite with irregular depression

laterally. Sixth somite (Figure 8e) highly sculptured

dorsally, with 4 pairs of longitudinal carinae, submedians armed, intermediates curving laterally,

each terminating in blunt lobe, laterals converging

with and fusing with marginals; median tubercle

and 6 rounded tubercles present lateral to submedian carina. Posterolateral angles with rounded projection. Submedian area of posterior margin lined

with rounded tubercles.

Telson (Figure Se,f) about 2 times broader than

long, with broad, inflated, longitudinally-sulcate

median boss, remainder of dorsal surface covered

with large, erect, rounded tubercles. 3 pairs of sharp

marginal teeth present, submedians with movable

apices. Denticles slender, sharp, small, 3-5, 4-7, 1.

Ventral surface smooth.

Uropod (Figure Se,g) broad, basal segment with

line of rounded tubercles extending to broad,

rounded distal lobe. Anterolateral margin strongly

convex, produced into broad, compressed, laterally

carinate lobe overhanging articulation of exopod.

Proximal segment of exopod shorter than distal,

outer margin completely lined with 9-10 blunt

spines, increasing in size distally, distal 3 slightly

recurved; dorsal surface with sharp tubercles. Distal

segment elongate-triangular, margin completely

lined with setae. Endopod curved mesially, margin

completely lined with setae, with line of 4 sharp

tubercles dorsally. Basal prolongation very short,

scarcely overreaching proximal segment of exopod,

produced into 2 blunt spines, inner longer.

MEASUREMENTS.—Only females examined, TL

11-13 mm. Borradaile (1898) reported a male and

2 females, 12-13 mm long. The female recorded by

Liu (1975) was 13.6 mm long. Other measurements

of female, TL 13 mm: carapace length 2.0; cornea

width 0.4; antennular peduncle length 2.0; rostral

plate length 0.2, width 0.4; fifth abdominal somite

width 2.2; telson length 1.0, width 2.0.

REMARKS.—Parvisquilla multituberailata differs

from the other member of the genus, P. sinuosa,

from Hawaii, primarily in having the telson ornamented with rounded tubercles rather than irregular carinae arranged in a scroll-like pattern.

The few specimens of this species examined are

all quite uniform, varying only in the numbers of

marginal denticles on the telson.

I can find no differences between the material of

P. multituberculata reported herein and the species

recently described from the Xisha Islands, P.

xishaensis Liu, 1975. Liu's species must be considered a synonym of P. multituberculata (Borradaile).

DISTRIBUTION.—Pacific Ocean, from Sandal Bay,

Lifu, Loyalty Islands (Borradaile, 1898; Kemp,

1913), Shi Island, Xisha Islands, Guangdong

[Kwangtung] Province, China (Liu, 1975), and now

from the Tonga Islands, Samoa and Moorea, Society Islands. Liu's specimen was taken on a coral

reef. The depth range for the species has not been

recorded.

Pullosquilla, new genus

DEFINITION.—Size very small, TL less than 30

mm. Body smooth, depressed, loosely-articulated.

Cornea (Figure 9a) subglobular. Rostral plate

(Figure 9a) triangular, unarmed anteriorly. Antennal protopod with ventral papilla. Carapace strongly

narrowed anteriorly, carinae and spines absent,

position of cervical groove indicated on lateral

plates only. Exposed thoracic somites lacking longitudinal carinae; eighth thoracic somite lacking

median ventral keel. Mandibular palp absent, 4 or

5 epipods present. Propodi of third and fourth

thoracic appendages broader than long, beaded or

ribbed ventrally. Claw (Figure 9b) stout, dactylus

with 10 or more teeth; propodus broad, superior

margin completely pectinate, with 4 movable spines

proximally; dorsal ridge of carpus undivided, terminating in spine; ischiomeral articulation terminal, ischium almost as long as merus, with strong

inferodistal spine on outer face. Endopods of

pereopods two-segmented, distal segment ovate or

subcircular on anterior 2 legs, slenderer on third.

Abdomen depressed, smooth, loosely-articulated,

anterolateral plates present; sixth somite unarmed

dorsally; articulated anterolateral plates present;

sixth somite unarmed posterolaterally, with curved,

ventrally-directed process anterior to articulation of

each uropod. Telson (Figure 9c, d) broader than

long, smooth dorsally, dorsal surface produced into
NUMBER 264

19

false eave overhanging marginal armature, obscuring most or all of marginal armature in dorsal view;

marginal armature including row of submedian

denticles, 1 movable submedian tooth, and 7 or

more (?) fixed lateral teeth and denticles. Uropod

(Figure 9c, e) flattened, proximal segment with distal, dorsal spine; proximal segment of uropodal

exopod shorter than distal, with slender, movable

spines on outer margin and stiff setae on inner, distal margin; endopod with strong proximal fold on

outer margin; spines of basal prolongation flattened,

triangular in cross-section, inner longer.

TYPE-SPECIES.—Austrosquilla litoralis Michel and

and Manning, 1971.

ETYMOLOGY.—The name is derived from the

Latin pullus, "young animal," in combination with

the name Squilla, alluding to the immature appearance of the species in the genus. The gender

is feminine.

REMARKS.—Pullosqiiilla includes three species,

one of which is described as new below; all three

species occur only in the Indo-West-Pacific region.

A key to the species is provided below, and additional records are given for P. litoralis.

Pullosquilla shares the armed ischium of the claw

with Austrosquilla Manning, 1966, and, because

of this feature, two of the species now assigned to

Pullosquilla had previously been placed in Austrosquilla. It seems likely that this feature developed

independently in these genera, for in most other

features Pullosquilla more closely resembles the

American genus Nannosquilla Manning, 1963.

These two genera share the small size as adults, the

loosely-articulated body, and having the telson being produced into a false eave which obscures the

marginal armature. Pullosquilla differs from Nannosquilla in having a triangular rather than a rectangular rostral plate, a papilla on the antennal

protopod, and a spine on the ischium of the claw.

The ischium and merus of the claw are much more

elongate in Pullosquilla than in Nannosquilla.

Key to Species of Pullosquilla

1. Sixth adbominal somite with 2 large ventral spines. Ventral surface of telson almost completely covered with spinules (Dactylus of claw with 12-15 teeth. 5 epipods present. Proximal segment of uropodal exopod with 3 outer spines and 2-4 inner stiff setae)

Pullosquilla thomassini, new species

Sixth abdominal somite unarmed ventrally. Ventral surface of telson unarmed 2

2. Dactylus of claw with 10-13 teeth. 5 epipods present. Proximal segment of uropodal exopod

with 3 outer spines and 2-5 inner stiff setae

Pullosquilla litoralis (Michel and Manning, 1971)

Dactylus of claw with 16-21 teeth. 4 epipods present. Proximal segment of uropodal exopod

with 4 outer spines and 3 inner stiff setae

Pullosquilla malayensis (Manning, 1968c)

Pullosquilla litoralis (Michel and Manning, 1971)

Austrosquilla litoralis Michel and Manning, 1971:237, fig. I.

Austrosquilla littoralis.—Thomassin, 1974:300, 301, 307.

MATERIAL.—Tiahura reef complex, Moorea, Society Islands;

B. Thomassin, leg., sta TIA-3; 1973: 1$, TL 13 mm; 29 .

TL 8.5-15 mm (SME). Data same; sta. TIA-19: 1 $ , TL 9

mm; 1 $, CL 2.5 mm (MNHNP). Data same; sta TIA-22:

1$, TL 19 mm (MNHNP). Data same; sta TIA-23: 2$,

TL 10-11 mm; 3 $ , TL 11-13 mm (USNM 169781). Aldabra

Island; J. Gamble: 13 , TL 15 mm; 2$ , TL 15-17 mm

(BMNH). Reef, Songoritelo, Madagascar; micro-atoll flat; B.

Thomassin. leg., sta 264; 26 Oct 1969: 1 9, TL 10.5 mm

(SME). Grand Recif, Tulear. Madagascar; B. Thomassin,

leg.; 1965: 1 £ , TL 13 mm; 3 9 • TL 13.5-15 mm; 2 juveniles,

TL 9.5 mm (1 Cl 1.8 mm) (in 6 lots; USNM 136294-136299)

(2g,2 9 SME).

REMARKS.—This species, originally described

from a single female taken on a sand beach in the

Marquesas Islands, is included here to document

its occurrence off Madagascar and Aldabra Island

in the western Indian Ocean, to demonstrate its

abundance there and at Moorea, and to record its

co-occurrence with another species of the genus,

described below, at the same station on the Tiahura

reef complex, Moorea. Both species occur on that

reef as well as in similar habitats on the Grand

Recif, Tulear, Madagascar.

The specimens reported here generally agree well

with the original description. There may be 10-13

teeth on the dactylus of the claw. The proximal

segment of the uropodal exopod has three movable
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spines laterally and two to five, usually three, stiff

setae on the inner distal angle. Some of the smaller

specimens, such as the female, TL 8.5 mm, from

Tiahura, probably are postlarvae, for the eave on

the telson is not yet formed and the submedian

denticles of the telson are marginal and visible in

dorsal view. At 10-11 mm the eave is poorly formed

but visible, and it attains the adult configuration

at a length of 13 mm.

This species apparently prefers sandy habitats in

relatively shallow water, on sand beaches or reef

flats. Thomassin (1974), the first to record its occurrence at Tulear, found it in coarse and medium

sands of the holes, couloirs, and patches of the

inner reef flat; he noted (p. 301) that they "reach

their greatest abundance in the inner part of the

moat, 250 m behind the boulder tract, generally

in scattered coral-growth flats." He also found the

species in the sandy bottom of a sea grass bed channel opening into the lagoon at Tulear.

DISTRIBUTION.—Nuka Hiva, Marquesas Islands;

Tiahura, Moorea, Society Islands; Songoritelo and

Tulear, Madagascar; and Aldabra Island. It occurs

in shallow water on sandy bottoms.

Pullosquilla thomassini, new species

FIGURE 9

MATERIAL.—Tiahura reef complex, Moorea, Society Islands;

B. Thomassin, leg., sta TIA-3; 1973; paratypes: 1^ , TL 10

mm (USNM 1.56276); 1 $ , TL 13 mm (SME). Tulear, Madagascar; sand flats; IIOE: no other data; paratype: 1$, TL

17 mm (USNM 156254). Grand Recif, lagoon enclave,

Grande Vasque (N zone), Tulear, Madagascar; 7-8 m; muddy

sand; B. Thomassin, leg., sta 773; 6 Jun 1972; paratype: 1 $ ,

CL 2.6 mm (MN'HNP). Plage de la Batterie, Tulear, Madagascar; quartzy sand; B. Thomassin, leg., sta 822; 22 Jun

1972; paratvpes: 1 9 , TL 18 mm (SME); 1 juvenile, TL 11

mm. 1 abdomen (USNM 156196). Internal creek, internal

slope, Grand Recif, Tulear, Madagascar; 7 m; coarse sand;

B. Thomassin, leg., sta 839; 7 Jul 1972: 1 $, TL 15 mm

(holotype; MNHNP); 1 $, TL 16 mm (paratype; USNM

156194).

DESCRIPTION.—Cornea (Figure 9a) set slightly

obliquely on stalk, overhanging stalk laterally. Eyes

reaching about to end of antennular peduncle.

Ocular scales fused into single, slender erect projection. Antennular peduncle slightly more than

1 15 carapace length. Antennal scale (Figure 9a)

about 1/4 carapace length; antennal protopod apparently lacking papillae. Rostral plate (Figure 9a)

triangular, almost cordiform, unarmed, apex depressed. Dactylus of claw (Figure 96) with 12-15

teeth. 5 epipods present. Basal segments of pereopods each with inconspicuous posterior spine. Ventral surface of sixth abdominal somite (Figure 9d)

with 2 strong spines on posterior margin. Telson

(Figure 9c, d) broader than long, subrectangular,

dorsal surface posteriorly produced into false eave

(shape of eave variable, often trilobed, with small,

rounded, median lobe), margin of eave with 12-22

small spines, fewer in smaller specimens; eave obscuring much of marginal armature in dorsal view;

marginal armature consisting of transverse row of

13-17 submedian denticles, and, lateral to these,

1 movable submedian tooth and 4 curved, fixed

lateral teeth, each flanked anteromesially by slender

denticle; midventral surface of telson (Figure 9d)

completely covered by numerous sharp fixed spines

and tubercles. Basal segment of uropod (Figure 9c)

with outer and dorsal carinae, dorsal carina terminating in 2 strong distal spines; proximal segment

of uropodal exopod (Figure 9c, e) much shorter

than distal, with 3 movable, spatulate spines, arranged in almost transverse row on distal margin;

inner distal angle of proximal segment of exopod

with 2-4 (usually 3) stiff setae; distal segment of

exopod and endopod very slender, curved mesially,

mesial margins strongly serrate for insertion of

setae; inner spine of basal prolongation of uropod

longer.

COLOR.—Faded in most specimens, but sometimes

marked with numerous dark chromatophores scattered over body. In some specimens dark pigment

concentrated laterally, occasionally with darker

spots on sixth thoracic, fifth and sixth abdominal

somites, and on telson.

MEASUREMENTS.—Males, TL 10-15 mm; females,

TL 13-18 mm. Other measurements of female, TL

17 mm: carapace length 3.1; cornea width 0.9; rostral plate length 1.3, width 1.2; antennal scale

length 0.8; fifth abdominal somite width 3.2; telson

length 1.7, width 3.0.

REIVTARKS.—This species with its ventrally spined

sixth abdominal somite, armed false eaves, and

spinulose ventral surface of the telson, can be distinguished easily from the other two species of the

genus, P. malayensis and P. litoralis. Like the latter

species, P. thomassini occurs both in Oceania and

in the western Indian Ocean, and the two species

have been taken together.
NUMBER 264 21

d

FIGURE 9.—Pullosquilla thomassini, new genus, new species, female paratype, TL 17 mm,

Tulear: a, anterior part of body; b, claw; c, sixth abdominal somite, telson, and uropod; d,

telson, ventral view; e, uropod, ventral view. (Setae omitted.)

ETYMOLOGY.—This species is named for B.

Thomassin, Station Marine d'Endoume, Marseille,

who collected most of the material reported above

and made it available to me for study.

DISTRIBUTION.—Tiahura, Moorea, Society Islands,

and localities around Tulear, Madagascar, on reef

flats in grass and sand. Shallow water to a depth

of 8 meters.

Family SQUILLIDAE Latreille, 1803

Genus A lima Leach, 1817

Alitna orientalis, new species

FIGURE 10

MATERIAL.—Takao, Formosa (Taiwan); Jun 1903; holotype:

1 $, TL 52 mm (ZSM). Mahe, Seychelles Islands; Mission

Zoologique; Jul-Sep 1966; paratype: 1 <J, TL 33.5 mm

(MRAC 52.236).

DESCRIPTION.—Size moderate, TL less than 60

mm. Surface of body rough. Eye (Figure 10a,6)

large, cornea stout, set almost transversely on stalk;

corneal indices 381-414. Rostral plate (Figure 10a)

as long as broad (juvenile) or broader than long

(adult), narrowed and with median carina anteriorly, apex broadly rounded. Anterolateral spines

of carapace strong, extending to base of rostral

plate. Median carina of carapace (Figure 10a) lacking distinct anterior bifurcation, intermediate carinae of carapace not extending to anterior margin.

Dactylus of claw with 5 teeth, outer margin of

dactylus flattened; dorsal ridge of carpus of claw

undivided, unarmed. Mandibular palp absent. 4

epipods present. Posterior 3 thoracic and abdominal somites with unarmed submedian and intermediate carinae. Lateral process of fifth thoracic

somite (Figure 10c) bilobed, sharp, anteriorlydirected anterior lobe set slightly below broadly

rounded posterior lobe. Lateral processes of sixth
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and seventh thoracic somites (Figure 10c) not

strongly bilobed, rounded laterally, process of sixth

somite with angled projection on anterior margin.

Anterior 5 abdominal somites with 8 carinae, sixth

somite with 6; abdominal carinae spined as follows: submedian 6, intermediate 5-6, lateral (4)

5-6, marginal (1-3) 4-5. Telson about as broad as

long, with low prelateral lobes, denticles 4, 7-9,

1, outer submedians (Figure lOd) and outer intermediates larger than remainder; ventral surface of

telson with long postanal keel. Uropod with 7-8

movable spines on outer margin of proximal segment of exopod, distalmost not extending to midlength of distal segment; 1 rounded lobe present

between spines of basal prolongation of uropod

(Figure lOe).

COLOR.—Largely faded in both specimens, but

posterior 3 thoracic and all abdominal somites with

dark posterior line, interrupted at intermediate and

lateral carinae on abdominal somites. Telson with

pair of large black spots anteriorly.

MEASUREMENTS.—Only male known, TL 33.5

mm; only female known, TL 52 mm. Other measurements of female holotype, TL 52 mm: carapace

length 12.0; cornea width 2.9; rostral plate length

1.1, width 1.8; fifth abdominal somite width 11.5;

telson length 8.5, width 9.3.

REMARKS.—Alima orientalis most closely resembles A. hieroglyphica (Kemp, 1911) (see Manning,

1969a: 135, fig. 40, for recent account), especially

in the broad eyes, carinate rostral plate, shape and

armature of the claw, and in having only one lobe

between the spines of the basal prolongation of

the uropod. The new species differs from A. hieroglyphica in having larger eyes, a more prominent

anterior projection on the lateral process of the

sixth abdominal somite, in lacking spines on the

submedian carinae of the fifth abdominal somite,

and in having rounded rather than triangular dark

patches on the telson.

The smaller paratype differs from the holotype

in having larger eyes, a narrower rostral plate,

slenderer lateral processes on the sixth and seventh

thoracic somites, and more spines on the abdominal

carinae. The first three of these differences can be

attributed to age. The spines of the abdominal

carinae of the holotype appear to have been worn

down.

FIGURE 10.—Alima orientalis, new species, female holotype. TL 52 mm, Formosa: a, anterior

part of body; b, eye; c, lateral processes of fifth, sixth, and seventh thoracic somites; d, submedian teeth of telson, ventral view (apices broken); e, basal prolongation of uropod, ventral

view. (Setae omitted.)
NUMBER 264 23

ETYMOLOGY.—The name is from the Latin orientalis, "of the east," alluding to the distribution of

the species.

DISTRIBUTION.—Indo-West-Pacific region, from

Formosa and the Seychelles Islands. The depth

range and habitat are unknown.

Busquilla, new genus

DEFINITION.—Eye (Figure \\a,b) very large, about

one-third carapace length, cornea bilobed, set

very obliquely on stalk. Inner margin of eyestalk longer than outer. Carapace with normal

complement of carinae, median carina lacking anterior bifurcation (Figure lla). Anterolateral angles

of carapace armed. Mandibular palp present. 4

epipods present. Dactylus of claw with 5 teeth.

Lateral processes of fifth, sixth, and seventh thoracic somites (Figure lie) bilobed, posterior lobe

much larger than anterior on both sixth and seventh somites. Abdomen with 8 carinae on anterior

5 somites, submedians low, indistinct anteriorly.

Telson flattened, lacking supplementary dorsal carinae, 3 pairs of marginal teeth present, submedians

with fixed apices (Figure 11/). Prelateral lobes

present. Basal prolongation of uropod (Figure llg)

unarmed on inner margin.

TYPE-SPECIES.—Busquilla plantei, new species.

ETYMOLOGY.—The name is derived with the

Latin prefix bu-, meaning "large," in combination

with the generic name Squilla, alluding to the

enormous eyes in representatives of this genus.

REMARKS.—Busquilla includes two species, one

of which, B. quadraticauda (Fukuda, 1911), previously was assigned to Oratosquilla Manning, 1968.

This new genus is clearly related to Oratosquilla,

differing in having much larger eyes, in lacking

the anterior bifurcation of the median carina of

the carapace, and in the structure of the lateral

processes of the fifth, sixth, and seventh thoracic

somites. These processes in Busquilla, although

similar to those of Oratosquilla, differ in that on

the fifth somite the two lobes are separate as well

as situated in different planes, and on the sixth

and seventh somites the anterior lobes are much

smaller than they are in corresponding species of

Oratosquilla. As in some species of Oratosquilla,

there are five rather than six teeth on the dactylus

of the claw.

Busquilla plantei, new species

FIGURE 11

MATERIAL.—Passe Lokobe, Madagascar; sand with Cerithiopsis; 17 m; R. Plante, leg.; May 1969; holotype: 1 $ , TL 53

mm (MNHNP). Grand Recif, Tulear, Madagascar; platform

between the buoys; B. Thomassin, leg., sta 631; 13 Apr 1972;

paratype: 1 $, TL 65.5 mm (USNM 156192).

DESCRIPTION.—Size moderate, TL to 65.5 mm.

Carapace and body smooth, even under magnification.

Eye (Figure \\a,b) very large, cornea bilobed,

set very obliquely on stalk. Eyes extending beyond

end of first segment of antennular peduncle. Ocular

scales subquadrate, inclined laterally, separated

mesially. Anterior margin of ophthalmic somite

with median spinule. Corneal indices 250-291.

Antennular peduncle longer than carapace. Dorsal processes of antennular somite produced into

anterolaterally-directed spines.

Rostral plate (Figure lla) slightly broader than

long, lateral margins converging on rounded apex.

Median carina absent.

Anterior width of carapace more than half median length. Anterolateral spines (Figure lla) very

short, not extending to base of rostral plate. Median carina almost completely absent anterior to

dorsal pit. Intermediate carinae parallel with laterals, not extending to anterior margin.

Dactylus of claw with 5 teeth, outer margin

sinuate, with low, obtuse projection basally. Dorsal ridge of carpus (Figure lid) irregular but

undivided, terminating in acute, rounded tooth.

Inferodistal angle on outer face of merus unarmed

(Figure lid).

Mandibular palp present. 4 epipods present.

Exposed thoracic somites lacking distinct submedian carinae, unarmed intermediate carinae

present on posterior 3 somites. Lateral process of

fifth somite (Figure lie) obscurely bilobed, lobes

separated by some distance, anterior lobe consisting of very small, anteroventrally-directed spine,

originating anterior to and ventral to minute,

laterally-directed posterior lobe. Lateral processes

of sixth and seventh somites bilobed (Figure lie),

posterior lobes larger, triangular, apices blunt.

Anterior 4 abdominal somites with very low,

indistinct submedian carinae, scarcely discernible

from medial longitudinal surface wrinkles; submedian carinae of fifth somite (Figure 11<?) distinct,
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

e

FIGURE 11.—Busquilla plantei, new genus, new species, male holotype, TL 53 mm, Madagascar:

a, anterior part of body; b, eye; c, lateral processes of fifth, sixth, and seventh thoracic somites

(insert shows enlargement of lateral process of fifth somite); d, carpus of claw; e, fifth abdominal somite; /, submedian teeth of telson, ventral view; g, basal prolongation of uropod, ventral

view. (Setae omitted.)

divergent. Abdominal carinate spined as follows:

submedian 5-6, intermediate (3)4-6, lateral (1)2-6,

marginal 1-5. Sixth somite with spinule ventrolaterally anterior to articulation of each uropod,

ventral surface with low median prominence on

posterior margin.

Telson flattened, length and width equal, with

3 pairs of marginal teeth. Prelateral lobes present,

shorter than margin of lateral teeth. Denticles

rounded, 3-4, 8-9, 1. Ventral surface with short

postanal keel.

Uropod broad, proximal segment of exopod

shorter than distal, with 8 movable spines on outer

margin, distalmost short, not extending to midlength of distal segment. Lobe on outer margin of

inner spine of basal prolongation of uropod (Figure

1 lg) large, rounded, margin concave.

COLOR.—Body covered with numerous small,

dark chromatophores. Posterior margin of posterior

3 thoracic and all 6 abdominal somites with dark

posterior line. Fifth abdominal somite with large

dark spot on posterior margin between lateral and

marginal carinae. Articular points between sixth

abdominal somite and telson dark. Posterior end

of median carina and bases of submedian teeth of

telson dark. Uropod with large black spot on exopod, covering distal half of proximal segment and

inner half of distal segment. Distal half of uropodal

endopod with few black chromatophores.

MEASUREMENTS.—Males only examined, TL

53-65.5 mm. Other measurements of male holotype, TL 53 mm: carapace length 11.0; anterior

width of carapace 6.8; cornea width 4.4; antennular

peduncle length 13.1; rostral plate length 2.0,

width 2.3; telson length 11.0, width 11.0.

REMARKS.—Busquilla plantei differs from B.

quadraticauda (Fukuda, 1911) in lacking a median

carina on the rostral plate, in having much more

divergent submedian carinae on the fifth abdominal somite, and in having fewer armed carinae on

the abdomen; the intermediates of the second

somite are unarmed in B. plantei. Kemp (1913, 55,

pi. 4: figs. 45-47) mentioned the median carina on

the rostral plate of his specimen of B. quadraticauda

but did not show it in his figure.

ETYMOLOGY.—This species is named for the collector of the holotype, R. Plante, Station Marine

d'Endoume, Marseille, France.

DISTRIBUTION.—Known only from two localities

off Madagascar.
NUMBER 264

25

Genus Clorida Eydoux and Souleyet, 1842

Clortda japonica, new species

FIGURE 12

MATERIAL.—Sanuki, Shikoku, Japan; 13 Mar 1897; holotype: lJ,TL44m m (ZMH K.7353).

DESCRIPTION.—Eye (Figure 12a, b) flask-shaped,

broad, cornea small, bilobed, much narrower than

adjacent stalk; basal half of mesial margins of eye

straight, subparallel; cornea width 1/3 eye length,

stalk width slightly more than y2 eye length. Rostral plate (Figure 12a) short, broader than long,

triangular, lacking median carina. Anterolateral

FIGURE 12.—Clorida japonica, new species, male holotype, TL 44 mm, Japan: a, anterior part

of body; b, eye; c, lateral processes of fifth, sixth, and seventh thoracic somites; d, carpus,

propodus, and dactylus of claw; e, abdomen; /, sixth abdominal somite, telson, and uropod;

g, submedian teeth of telson, ventral view; h, basal prolongation of uropod, ventral view

(rounded lobe between uropod spines foreshortened in this view; it curves dorsally). (Setae

omitted.)
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

margins of carapace slightly sinuous, anterolateral

angles armed. Mandibular palp present. 4 epipods

present. Dactyl us of claw (Figure \2d) with 4 teeth,

proximal with indication of fifth tooth at base.

Lateral process of fifth thoracic somite (Figure 12c)

consisting of blunt, laterally directed spine, ventral

spine absent. Lateral processes of sixth and seventh

thoracic somites (Figure 12c) rounded anterolaterally and posterolaterally, process of seventh somite

more truncate. Thoracic somites lacking submedian

carinae. Anterior 4 abdominal somites lacking submedian carinae, fifth somite with low, unarmed

submedian carinae (Figure 12e). Abdominal carinae

spined as follows: submedian 6, intermediate 5-6,

lateral 5-6, marginal 5. Sixth abdominal somite

(Figure 12/) lacking supplementary spinules on

posterior margin, dorsal surface rugose lateral to

submedian carinae. Telson (Figure 12/) with prelateral lobes well-marked dorsally, dorsal ornamentation as illustrated; low, long postanal keel present

ventrally. Marginal denticles of telson sharp, 3-4,

7, I, outer margin of intermediate tooth tuberculate. Uropodal exopod (Figure 12/) with 6-7 movable spines on outer margin of proximal segment,

distalmost slender, not extending to midlength of

distal segment. Basal prolongation of uropod (Figure 12h) with 7-8 slender fixed spines on inner

margin, lobe between apical spines small, rounded.

COLOR.—Largely faded in holotype, but rostral

plate dark; carapace with diffuse patch of dark pigment medially, margin outlined in black pigment;

each body segment with dark posterior line.

MEASUREMENTS.—Only known specimen, male

holotype, TL 44 mm. Other measurements: carapace length 8.6; cornea width 0.8; eye length 2.4;

eyestalk width 1.3; rostral plate length 1.1, width

1.7; antennular peduncle length 8.3; fifth abdominal somite width 10.8; telson length 6.8, width 9.8.

REMARKS.—Clorida japonica belongs to the section of the genus that includes species with a mandibular palp and traces of submedian carinae on

at least some of the anterior five abdominal somites; it shares the latter feature with C. verrucosa

(Hansen, 1926) and C. merguiensis (Tiwari and

Biswas, 1952) (see Manning, 1976b for recent accounts of these species.) It differs from both of

them in having a much broader eye, a much shorter

rostral plate, fewer spines on the abdominal carinae (none of the carinae of the anterior four somites are armed), and in lacking ventral spines on

the fifth abdominal somite. It also resembles C.

depressa (Miers, 1880), but that species has all five

marginal carinae of the abdomen armed and lacks

submedian carinae on the anterior five abdominal

somites as well as a postanal keel.

ETYMOLOGY.—The name is derived from the

type-locality in Japan.

DISTRIBUTION.—Known only from the type-locality, Sanuki, Shikoku Island, Japan (probably Sanuki Province, now Kagawa Prefecture).

Clorida latispina Manning, 1968

Squilla latreillii.—Gravier, 1930:524 [not C. latreillei Eydoux

and Souleyet, 1842].

Clorida latispina Manning, 1968c:247, fig. 3.

MATERIAL.—Cap St. Jacques, Tonkin, Viet Nam; 30 Sep

1926: 1 $ , TL 67 mm (MNHNP).

REMARKS.—This specimen was identified by

Gravier (1930) as C. latreillei. It agrees in most respects with the original account of C. latispina,

differing only in having four rather than two or

three submedian denticles on the telson. As in the

types of C. latispina, submedian carinae are not

present on the thoracic somites anterior to the

eighth.

DISTRIBUTION.—Indo-Malayan area, from off

Hong Kong in 34 fms (62 m), Tonkin, and Khota

Bharu, Malaysia.

Clorida malaccensis Manning, 1968

Clorida malaccensis Manning, 1968c:244, fig. 2.

Clorida malaccensis var. moluccensis Moosa, 1973:4 [listed],

19, fig. 3.

MATERIAL.—Nosy Mitsio, Madagascar; 42 m; muddy sand;

R. Plante, leg.; 22 Jun 1970: 1 specimen, fragmented (SME).

REMARKS.—This single specimen, represented by

the anterior half of the carapace and a claw, is

clearly identifiable with C. malaccensis, previously

known from two localities in Indo-Malaya. There

are five teeth on the claw, and all features present,

including the color pattern, agree with the holotype.

I believe that Moosa's subspecies should be synonymized with C. malaccensis. His material is said

to have four rather than five teeth on the claw,
NUMBER 264 27

although the proximal small tooth which he stated

(p. 21) to be absent is shown in his fig. 3c. The

ocular scales in his specimens are slightly different

in shape, but there appear to be no other differences.

DISTRIBUTION.—Indo-West-Pacific region, from

the Strait of Malacca, the type-locality (Manning,

1968c); from off Aru, Indonesia (05°37'S, 134°10'E),

in 55-66 m on mud and fine shelly grit, and the

Arafura Sea (04°52'S, 135°25'E), in 90 m on mud

and sand (both Moosa, 1973); and now from Madagascar, in 42 m on muddy sand.

Clorida minor (Jurich, 1904)

FIGURE 13

Squilla minor Jurich, 1904:364, pi. 25; figs. 4-4a [p. 6, pi. 1:

figs. 4—4a on separate].—Kemp, 1913:40 [discussion].—Manning, 1968b: 131 [listed; transferred to Squilloides].

Squilloides minor.—Manning and Serene, 1968:118 [listed].

Clorida fallax.—Manning, 1970:1430 [not Clorida fallax (Bouvier, 1914)].

MATERIAL.—Zanzibar; Valdivia sta 244; 50 m; syntypes: 1 $,

TL ca. 20 mm; 1 9, TL ca. 16 mm (ZMB). Madagascar;

23°21'50"S, 43°38'20"E; 10 m; muddy sand; R. Derijard, leg.,

sta D-2; 14 Aug 1963: 1$. TL 23 mm (USNM 127457).

FIGURE 13.—Clorida minor (Jurich, 1904), male syntype, TL ca. 20 mm, off Zanzibar: a, anterior part of body; b, lateral processes of fifth, sixth, and seventh thoracic somites; c, claw

(tip of dactylus broken or deformed); d, sixth abdominal somite, telson, and uropod; e, uropod,

ventral view. (Setae omitted.)
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Madagascar; 23O21'56"S, 43°38'09"E; 14 m; sandy mud; R.

Derijard, leg., sta D-55; 28 Sep 1963: 1 9, CL 4.9 mm

(MNHNP). Nosy Mitsio, Madagascar; 40 m; sand; R. Plante,

leg.: 1 $ , TL 16.5 mm (SME).

DESCRIPTION.—Eye (Figure 13a) short and stout,

cornea bilobed, broader than and set obliquely on

stalk; mesial margin of eye sinuous; cornea width

about 2/s eye length, stalk width more than half

but less than 2/3 eye length. Rostral plate (Figure

13a) short, length and width subequal, lacking

median carina. Anterolateral margins of carapace

(Figure 13a) straight or slightly concave, anterolateral angles armed. Mandibular palp absent. 4

epipods present. Dactylus of claw (Figure 13c) slender, with 6 teeth. Lateral process of fifth thoracic

somite (Figure 136) consisting of short, anterolaterally-directed spine; prominent ventral spine also

present. Lateral processes of sixth and seventh thoracic somites (Figure 136) broadly rounded posterolaterally. Posterior 3 thoracic and anterior 5

abdominal somites lacking submedian carinae. Abdominal carinae spined as follows: submedian 6,

intermediate 6, lateral 6, marginal 5; posterolateral

margin of fifth somite angled, not broadly rounded

as in C. fallax, usually armed with minute spinule.

Sixth abdominal somite (Figure 13d) lacking supplementary spinules on posterior margin, dorsal

surface, lateral to submedian carinae, rugose. Telson (Figure 13d) lacking prelateral lobes laterally

(lobes formed dorsally, not distinct laterally), dorsal

ornamentation as illustrated; accessory median carinae of telson reduced to line of small tubercles,

scarcely visible in some specimens. Ventral surface

of telson with short postanal keel. Marginal denticles of telson sharp, 3^1, 7-8, 1. Uropodal exopod

(Figure 13d, e) with 6-8 movable spines on outer

margin of proximal segment, distal 2 spatulate,

distalmost not extending to midlength of distal

segment. Basal prolongation of uropod (Figure 13e)

with 6-9 slender fixed spinules on inner margin

and 2 broad, rounded lobes between apical spines,

inner much the larger.

COLOR.—Carapace with dark pigment anteriorly,

U-shaped dark patch near midlength, and broader,

darker band across posterior third. Merus of claw

with dark patch dorsally. Exposed thoracic and

abdominal somites with squarish dark patch on

midline. Fifth thoracic somite with dark spot on

lateral process, remainder of thoracic and abdominal somites dark laterally. Telson with dark patch

along median carina and at bases of intermediate

and lateral marginal teeth. Distal segment of exopod and distal y3 of endopod black.

MEASUREMENTS.—Males, TL 16.5-23 mm; only

intact female, TL ca. 16 mm. Other measurements

of male, TL 23 mm: carapace length 4.6; cornea

width 1.2; eye length 1.5; eyestalk width 0.9; rostral plate length 1.0, width 0.7; fifth abdominal

somite width 5.0; telson length 3.1, width 4.0.

REMARKS.—Clorida minor, which was assigned

by me (1968b) to Squilloides on the basis of Jurich's

incomplete account, is a distinctive species of Clorida that closely resembles C. fallax (Bouvier, 1914)

(see Manning, 1968a:8, fig. 2). Clorida minor differs from C. fallax in numerous features: the claw

is slenderer and the dactylus is armed with six

rather than four teeth; the posterolateral angle of

the fifth abdominal somite is angled and usually

spined rather than broadly rounded; the accessory

median carinae of the telson are formed by lines

of tubercles; the ventral surface of the telson is

ornamented with a postanal keel only; and there

are six to eight rather than five movable spines

on the uropodal exopod.

DISTRIBUTION.—Western Indian Ocean, from

Zanzibar and Madagascar, in depths between 10

and 50 m.

Genus Cloridopsis Manning, 1968

Cloridopsis aquilonaris, new species

FIGURE 14

Squilla scorpio.—Balss, 1910:8 [part] [not Cloridopsis scorpio

(Latreille, 1828)].

Chloridella scorpio.—Schmitt, 1931:133 [part] [not Cloridopsis scorpio (Latreille, 1828)].

MATERIAL.—Okayama Sea, Japan; Doflein, leg., no. 2389;

1904-1905; paratypes: 2$, TL 66.5-84 mm; 1 9, TL 69.5 mm

(ZSM). Mui Hua, Foochow, China; A. S. Pearse, leg.; Apr

1930; holotype: 1 ? , TL 88 mm (USNM 168778). Same data;

paratypes: 7 $, TL 62-93 mm; 6$ , TL 46-89 mm (USNM

120326). Foochow, China; R. Kellogg, leg.; 1924; paratypes:

1 $, TL 78 mm; 19 TL 77 mm (USNM 58745).

DIAGNOSIS.—Size moderate, total length of adults

less than 100 mm. Eye (Figure 14a, b) slender, stalk

width about i/2 eye length, inner margin of stalk

not expanded; cornea broader than stalk. Corneal

indices 559-707. Rostral plate (Figure 14a) short,

as broad as or broader than long, with median

carina. Mandibular palp present. 2 epipods pres-
NUMBER 264 29

FIGURE 14.—Cloridopsis aquilonaris, new species, male paratype, TL 84 mm, Okayama Sea:

a, anterior part of body; b, eye; c, lateral processes of fifth, sixth, and seventh thoracic somites;

d, carpus of claw; e, submedian teeth of telson, ventral view; /, basal prolongation of uropod,

ventral view. (Setae omitted.)

ent. Dactylus of claw with 5-6 (usually 5) teeth.

Lateral process of fifth thoracic somite (Figure 14c)

in form of broad, blunt spine, slightly inclined

ventrally, directed anterolaterally, with prominent

black spot basally. Abdominal carinae spined as

follows: submedian 6, intermediate 5-6, lateral

5-6, marginal 3-5. Telson denticles blunt (Figure

\Ae), 1-3, 3-5, 1. Ventral surface of telson lacking

postanal keel. Uropod with 6-7 spines on proximal

segment of exopod, distalmost short.

COLOR.—Posterior margin of carapace, posterior

3 thoracic somites, and all abdominal somites with

dark posterior line. Lateral process of fifth thoracic

somite (Figure 14c) with prominent black spot at

base. Second abdominal somite with dark, rectangular patch medially. Bases of teeth of telson dark.

Uropodal exopod with dark patch distally on

proximal segment.

MEASUREMENTS.—Males, TL 62-93 mm; females,

TL 46-89 mm. Other measurements of male holotype, TL 88 mm: carapace length 18.8; cornea

width 2.9; rostral plate length 1.8, width 1.8; fifth

abdominal somite width 21.0; telson length 16.5;

width 18.5.

REMARKS.—Cloridopsis aquilonaris is very similar

to both C. scorpio (Latreille, 1828) and C. immaculata (Kemp, 1913) (see Kemp, 1913, for accounts of

these species). It resembles C. immaculata and differs from C. scropio in having a short rostral plate,

and it resembles C. scorpio and differs from C. immaculata in having a prominent black spot at the

base of the lateral process of the fifth thoracic somite. It further differs from C. scorpio in that

the lateral process of the fifth thoracic somite is

blumer, deflected ventrally, and directed anterolaterally rather than anteriorly. As in other species,

in males the chelae are heavier and the carinae of

the sixth abdominal somite and telson are noticeably inflated.

Cloridopsis aquilonaris is the northern counterpart of C. scorpio. The specimens from Japan identified as C. scorpio by Balss (1910) proved to be

C. aquilonaris, as did part of the material from

China identified as C. scorpio by Schmitt (1931).

Both species occur off China. It seems likely that

some or all records of C. scorpio from Japan

(Komai, 1927) may be referable to C. aquilonaris.

ETYMOLOGY.—The name is from the Latin aquilonaris, "northern," alluding to the distribution of

the species in relation to C. scorpio.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—Known only from China and

Japan.

Genus Harpiosquilla Holthuis, 1964

Harpiosquilla malagasiensis, new species

FIGURE 15

MATERIAL.—Tamatave, Madagascar; von Rosenberg leg.;

holotype: 1 $ , TL 196 mm (ZMH K.7364).

DESCRIPTION.—Size large, total length of adult

almost 200 mm. Cornea (Figure 15a, b) large, corneal index 327 in holotype. Rostral plate (Figure

15a, c) short, length and width subequal, apex

rounded, lacking apical projection. Carapace with

median carina. Dactylus of claw (Figure \be) with

8 teeth; opposable margin of propodus of claw with

1-2 smaller spines and several minute denticles

between largest spines. Fifth thoracic somite (Figure 15d) rounded laterally, irregular, unarmed.

Posterior 3 thoracic somites with submedian and

intermediate carinae, none armed. Ventral keel of

eighth thoracic somite broadly rounded. Anterior

3 abdominal somites with distinct submedian carinae, submedians very low, scarcely discernible, on

fourth and fifth somites. Abdominal carinae spined

as follows: submedian 6, intermediate 2-6, lateral

1-6, marginal 1—5. Telson denticles 4-5, 11, 1;

marginal carina of telson (Figure 15/) about 3

times as long as lateral carina. Postanal keel of

telson extending about % distance from anal pore

to posterior margin. Uropodal exopod (Figure 15/)

with 9 movable spines on outer margin of proximal

segment.

COLOR.—Completely faded.

MEASUREMENTS.—Only known specimen, female

holotype, TL 196 mm. Other measurements: carapace length 38.3; cornea width 11.7; rostral plate

length 5.9, width 6.0; raptorial propodus length

51.2; telson length 33.1, width 34.1.

REMARKS.—This new species belongs to the section of Harpiosquilla which includes those species

with a short rostral plate lacking a distinct anterior projection: H. annandalei (Kemp, 1911), H.

japonica Manning, 1969c, H. stephensoni Manning,

1969c, and H. intermedia Manning and Michel,

1973. (See Manning, 1969c; Manning and Michel,

1973). Harpiosquilla malagasiensis most closely resembles H. intermedia, but it differs from it in

having a much shorter rostral plate, less prominent

submedian carinae on the fifth abdominal somite,

and a comparatively longer marginal carina on the

telson. These differences are not known to change

with age or sex.

ETYMOLOGY.—The specific name is derived from

the type-locality, Madagascar (Malagasy Republic).

DISTRIBUTION.—Known only from the typelocality.

Tuleariosquilla, new genus

DEFINITION.—Size very small, adults TL 15 mm

or less. Eye (Figure 16a, b) large, cornea strongly

bilobed, noticeably broader than stalk. Inner margin of eye slightly longer than outer. Ocular scales

(Figure 16a) low, separate. Antennular somite (Figure 16a) elongate. Carinae of carapace reduced,

median and intermediate carinae absent. Anterolateral angles of carapace (Figure 16a) unarmed,

lateral plates very small, gastric grooves scarcely

indicated. Mandibular palp absent. 3 epipods present. Dactylus of claw (Figure 16d) with 4 teeth.

Lateral processes of fifth, sixth, and seventh thoracic somites (Figure 16c) single. Anterior 5 abdominal somites lacking submedian carinae, sixth

somite (Figure 16e) only with submedian carinae.

Telson (Figure 16e) flattened, dorsal carinae reduced; 3 pairs of marginal teeth present, submedian

with movable apices. Prelateral lobes absent. Inner

margin of basal prolongation of uropod (Figure

16/) smooth, unarmed.

TYPE-SPECIES.—Tuleariosquilla parvula, new

species.

ETYMOLOGY.—The generic name has been derived from a combination of the type-locality, Tulear, with the generic name Squilla. The gender is

feminine.

REMARKS.—Tuleariosquilla is based on two specimens of an unusually small squillid which appears

to be fully mature at a length of 13.5—14 mm. As

in Leptosquilla Miers, 1880, Tuleariosquilla has

an elongated antennular somite which is not overreached by the rostral plate and the mandibular

palp has been suppressed. Tuleariosquilla differs

from Leptosquilla in numerous features, the most

important of which are: the eye is very large and

the cornea is bilobed, the ocular scales are separate,

there are only 4 teeth on the claw and 3 epipods,

and the inner margin of the basal prolongation of
NUMBER 264 31

FIGURE 15.—Harpiosquilla malagasiensis, new species, female holotype, TL 196 mm, Madagascar: a, anterior part of body; b, eye; c, rostral plate; d, lateral processes of fifth, sixth,

and seventh thoracic somites; e, propodus and dactylus of claw; /, uropod. (Setae omitted.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the uropod is completely smooth. I suspect that the

similarities between these genera are superficial and

that they are not closely related.

The elongated antennular somite has appeared,

presumably independently, in four stomatopod

genera representing three families: Eurysquilloides

Manning, 1963 (Family Gonodactylidae), Parvisquilla Manning, 1973 (Family Lysiosquillidae), and

Leptosquilla Miers, 1880 and Tuleariosquilla, new

genus (Family Squillidae). Its significance is unknown.

Tuleariosquilla parvula, new species

FIGURE 16

MATERIAL.—Songoritelo Reef, near Tulear, Madagascar;

micro-atoll platform; B. Thomassin, leg., sta 265; 26 Oct

1969: 1 $, TL 13.5 mm (holotype, MNHNP); 1 $, TL 14 mm

(paratype, USNM 156191).

DESCRIPTION.—Eye (Figure 16a, b) large, cornea

strongly bilobed, set almost transversely on stalk.

Eyes not extending to end of first segment of antennular peduncle. Ocular scales (Figure 16a) low,

separate. Anterior margin of ophthalmic somite

unarmed, emarginate. Corneal index 338.

Antennular somite (Figure 16a) elongate, enlarged, almost as wide as anterior margin of carapace. Dorsal processes of antennular somite reduced

to acute anterolateral angles of somite. Antennular

peduncle almost as long as carapace.

Antennal protopod elongate, extending anteriorly

to or beyond antennular somite. Antennal peduncle

almost overreaching eye. Antennal scale very small.

Rostral plate (Figure 16a) triangular, slightly

broader than long, rounded anteriorly, lacking

carinae or spines, not extending anteriorly to midlength of antennular somite.

Carapace short, strongly narrowed anteriorly,

lateral plates not covering chelipeds in dorsal view.

Carinae reduced, traces of reflected marginals

scarcely visible posteriorly; no other carinae visible.

Gastric grooves very faint, cervical groove absent.

Anterolateral angles (Figure 16a) unarmed.

FIGURE 16.—Tuleariosquilla parvula, new genus, new species, male holotype, TL 13.5 mm,

Tulear: a, anterior part of body; b, eye; c, lateral processes of fifth, sixth, and seventh thoracic

somites; d, carpus, propodus, and dactylus of claw; e, sixth abdominal somite, telson, and

uropod; /. basal prolongation of uropod, ventral view. (Setae omitted.)
NUMBER 264 33

Claw (Figure 16d) large, merus very stout, completely visible in dorsal view. Dactylus with 4 teeth,

outer margin flattened, with prominent basal notch.

Propodus deep, superior margin fully pectinate.

Dorsal ridge of carpus undivided, terminating in

obtuse blunt lobe.

Exposed thoracic somites (Figure 16c) lacking

longitudinal carinae. Lateral process of fifth thoracic somite forming rounded, spatulate lobe; blunt

ventral spin present on each side beneath lateral

process. Lateral processes of sixth and seventh thoracic somites rounded laterally, pointed posteriorly,

unarmed. Ventral keel of eighth thoracic somite inconspicuous. Male copulatory tubes well-developed.

Anterior 5 abdominal somites lacking submedian

carinae, intermediates apparently absent also, low

unarmed lateral carinae present on each somite.

Fifth somite bluntly pointed posterolaterally. Sixth

somite (Figure 16<?) with 3 pairs of spines dorsally,

submedians and laterals with low carinae.

Telson (Figure 16e) broader than long, median

carina sharp, terminating in slender spine. Submedian teeth only with short, inconspicuous dorsal

carina, remainder of surface smooth. Denticles

sharp, 6-7, 6-7, 0. Ventral surface of telson smooth.

Proximal segment of uropodal exopod (Figure

shorter than distal, with 5-6 sharp, movable

spines on outer margin, distalmost extending almost to end of distal segment. Basal prolongation

of uropod (Figure 16/) produced into 2 spines,

inner much longer, with broad, rounded lobe on

outer margin; inner margin of basal prolongation

smooth, with no trace of serrations or tubercles.

COLOR.—Almost completely faded, but with traces

of large brown and smaller white chromatophores

in longitudinal row on each side of abdomen. Apex

of median carina and posterior margin of telson

darker than surface.

MEASUREMENTS.—Males only known, TL 13.5-14

mm. Other measurements of male holotype: carapace length 2.7; cornea width 0.8; antennular peduncle length 2.6; rostral plate length 0.4, width

0.3; telson length 1.6, width 2.0.

REMARKS.—Although initially I identified these

specimens as postlarval squillids, closer examination revealed the presence of well-developed copulatory tubes, indicating that they are fully mature.

The combination of characters is so unique that

T. parvula should be recognized without difficulty.

DISTRIBUTION.—Known only from the type-locality, Songoritelo reef, near Tulear, Madagascar.
Literature Cited

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1910. Ostasiatische Stomatopoden: Beitrage zur Naturgeschichte Ostasiens, Herausgegeben von Dr. F.

Doflein. Abhandlungen der Bayerischen Akadetnie

der Wissenschaften, supplement 2, 2:1-11, figures

1-2.

Barnard, K. H.

1958. Further Additions to the Crustacean Fauna-List of

Portugese East Africa. Mem&rias do Museu Dr.

Alvaro de Castro, 4:5-23, figures 1-7.

Berthold, A. A.

1827. Latreille's Naturliche Familien des Thierreichs, aus

dent Franzosischen mit Anmerkungen und Zusdtzen.

x + 606 pages. Weimar.

Borradaile, L. A.

1898. Stomatopoda. On Some Crustaceans from the South

Pacific, Part 1. Proceedings of the Zoological Society

of London, 1898:32-38, plates 5-6.

1899. On the Stomatopoda and Macrura Brought by Dr.

Willey from the South Seas. In Willey, Zoological

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36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Manning, Raymond B., and Alain Michel

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