Synopses of the Indo-West-Pacific

Species of Lysiosquilla Dana, 1852

(Crustacea: Stomatopoda:

Lysiosquillidae)

RAYMOND B. MANNING

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 259
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 25 9

Synopses of the Indo-West-Pacific

Species of Lysiosquilla Dana, 1852

(Crustacea: Stomatopoda: Lysiosquillidae)

Raymond B. Manning

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978
ABSTRACT

Manning, Raymond B. Synopses of the Indo-West-Pacific Species of Lysiosquilla Dana, 1852 (Crustacea: Stomatopoda: Lysiosquillidae). Smithsonian Contributions to Zoology, number 259, 16 pages, 13 figures, 1978.—The five species

of Lysiosquilla from the Indo-West-Pacific region are characterized. Two previously known species, L. maculala (Fabricius) and L. sulciroslris Kemp, and a

new species, L. sulcata, are described and illustrated. Comparative diagnoses and

sketches are provided for these species as well as for L. capensis Hansen and

L. tredecimdentata Holthuis.

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

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrca

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Manning, Raymond B. 1934-

Synopses of the Indo-West-Pacific species of Lysiosquilla Dana, 1852.

(Smithsonian contributions to zoology ; no. 259)

Bibliography: p.

1. Lysiosquilla. 2- Crustacea—Classification. 3. Crustacea—Indian Ocean. 4. Crustacea—

Pacific Ocean. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to

zoology ; no. 259.

QL1.S54 no. 259 [QL444.M375] 591'OSs [595'.382] 77-10934
Contents

Page

Introduction 1

Key to Indo-West-Pacific Species of Lysiosquilla 2

Lysiosquilla capensis Hansen, 1895 3

Lysiosquilla maculata (Fabricius, 1793) 3

Lysiosquilla sulcata, new species 7

Lysiosquilla sulcirostris Kemp, 1913 11

Lysiosquilla tredecimdentata Holthuis, 1941 13

Literature Cited 16

i l l

Synopses of the Indo-West-Pacific

Species of Lysiosquilla Dana, 1852

(Crustacea: Stomatopoda: Lysiosquillidae)

Raymond B. Manning

Introduction

The pantropical stomatopod genus Lysiosquilla,

which includes the largest known stomatopods, now

comprises 12 species, five of which occur only in

the Indo-West-Pacific region. Two of the five, L.

capensis Hansen and L. tredecimdentata Holthuis,

have been redescribed in recent papers (Manning,

1969 and 1968, respectively). Two other previously

known species, L. maculata (Fabricius) and L. sulcirostris Kemp, are redescribed below, and the fifth

is named below.

Lysiosquilla maculata, the largest and the most

common species in the Indo-West-Pacific region,

has never been characterized adequately. The most

complete account, that by Kemp (1913), was based

on specimens of both L. maculata and L. tredecimdentata. It seems likely that many of the references

to L. maculata, summarized in Holthuis (1967),

apply to one or more of the other species reported

herein. For this reason the ranges of most of the

species included in this study cannot be determined

with any precision and all references to L. maculata

in the literature require verification. The synonymies compiled by Holthuis (1967) have not been

duplicated here, but are referred to along with

references to the original description and other

selected ones.

Differential diagnoses and stylized comparative

Raymond B. Manning, Department of Invertebrate Zoology,

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

outline drawings (Figures 9-13) of diagnostic features of the five Indo-West-Pacific species of Lysiosquilla are included, supplementing the more

complete accounts of L. maculata, L. sulcirostris,

and L. sulcata, new species. The stylized outline

drawings, patterned after those introduced by

Hobbs (1974) for crayfishes, should simplify identification of these members of the Indo-West-Pacific

stomatopod fauna.

Several simple diagnostic characters for Lysiosquilla apparently have not been used in the past.

The shape and pigmentation of the antennal scale

can be determined readily: it is slender (Figures

lla, 12a, 13a) and outlined by dark pigment or

with the surface covered with dark chromatophores in L. capensis, L. sulcirostris, and L. tredecimdentata, broadly oval (Figures 9a, 10a) with

an irregular surface spot in L. maculata and L.

sulcata, new species. In the three species with a

slender antennal scale, there is a sharp, fixed anterior projection on the protopod above the articulation of the antennal peduncle (Figures lla, 12a,

13a); those species with a broad antennal scale

lack this projection. The fixed projection should

not be confused with the supple antennal papillae

found in all members of this genus. Finally, the

presence of dark pigment on the apex of the uropodal endopod has proved to be a reliable specific

character.

Sexual dimorphism, affecting primarily the size

and number of teeth on the claw of adult females,

has been documented for two of the Indo-West1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pacific species and may well occur in all species.

Kemp (1913) and Bigelow (1931), among others,

have reported on dimorphism in L. maculata, and

Serene (1954) noted it on L. sulcirostris. In adult

females the claw is smaller than in males, the size

and number of teeth on the claw are reduced, and

the carpus and propodus may be ornamented with

numerous simple setae.

Accounts of the postlarval forms of L. maculata

and L. sulcirostris from India were published by

Alikunhi (1967). As noted below under the account

of L. tredecimdentata, Alikunhi may have been

dealing with that species rather than L. maculata.

Furthermore, it is not certain from his account

whether his observations were based upon L. sulcirostris or upon L. sulcata, both of which, as

adults, have less than 10 teeth on the dactylus of

the claw. Michel (1970) recorded the occurrence

of larvae and postlarvae of three distinct species

of Lysiosquilla from the central Pacific, one similar

to L. capensis (not known outside of South African

waters), one like L. maculata, and one resembling

L. sulcirostris. It seems likely that additional species of Lysiosquilla, possibly from deeper water,

will be found in the future in the Indo-WestPacific region.

Measurements and terms used in the accounts

have been explained in earlier papers (Manning,

1968, 1969). All measurements are in millimeters

(mm). Total length (TL) is measured on the midline from the anterior margin of the rostral plate

to the posterior median margin of the telson.

The following abbreviations are used to designate repositories:

BMNH British Museum (Natural History), London

MNHNP Museum National d'Histoire Naturelle, Paris

NMV Naturhistorisches Museum, Vienna

RMNH Rijksmuseum van Natuurlijke Historic Leiden

SMF Natur-Museum und Forchungsinstitut

Senckenberg, Frankfurt am Main

USNM National Museum of Natural History,

Smithsonian Institution, Washington (formerly

United States National Museum)

ZMB Zoologisches Museum an der HumboldtUniversitat zu Berlin, Berlin

ZSI Zoological Survey of India, Calcutta

ACKNOWLEDGMENTS.—I thank B. Campbell,

Queensland Museum, Brisbane, for lending the

holotype of L. miersii De Vis; J. Forest, Museum

National d'Histoire Naturelle, Paris, for providing

working space there in 1971 and for the subsequent

loan of material; H.-E. Gruner and G. Hartmann,

Zoologisches Museum, Berlin, for providing working space in 1971; L. B. Holthuis, Rijksmuseum

van Natuurlijke Historic Leiden, for many kindnesses, including numerous loans, providing working space on several visits, and for discussions; R.

W. Ingle, British Museum (Natural History), for

loans and for arranging accommodations and access

to the collections there on several visits; G. Pretzmann, Naturhistorisches Museum, Vienna, for

loans and for arranging accommodations and access

to the collections in 1971; K. K. Tiwari, Zoological

Survey of India, for his help during a visit to Calcutta in 1972; M. Turkay and the late R. Bott,

Natur-Museum und Forschungsinstitut Senckenberg, for loans and for arranging accommodations

and working space in 1971; and Torben Wolff,

Universitetets Zoologiske Museum, who provided

working space in 1971.

Part of this study, including visits to European

Museums in 1971, was supported by the Smithsonian Institution through its Research Awards Program. The visit to the Zoological Survey of India

in 1972 was supported by the Foreign Currency

Program of the Smithsonian. All of the illustrations

were prepared by my wife Lilly.

I thank Cynthia Hemming and Anne Cohen for

reading a rough draft of the manuscript and for

checking references. Horton H. Hobbs, Jr., critically reviewed the manuscript.

Key to Indo-West-Pacific Species of Lysiosquilla

1. Antennal potopod with angled dorsal projection. Antennal scale slender, length 3 or more

times greatest width. Ventral keel of eighth thoracic somite sharp (apex of uropodal

endopod dark) 2

Antennal protopod lacking angled dorsal projection. Antennal scale broad, length less than

3 times greatest width. Ventral keel of eighth thoracic somite rounded 4

2. Rostral plate lacking median carina. Dactylus of claw with 15-17 teeth

L. capensis Hansen, 1895

Rostral plate with median carina. Dactylus of claw with less than 15 teeth 3
NUMBER 259

Key to Indo-West-Pacific Species of Lysiosquilla (Con't)

3. Median carina on rostral plate flanked laterally by longitudinal grooves. Dactylus of claw

with 7-8 teeth L. sulcirostris Kemp, 1913

Median carina on rostral plate not flanked laterally by longitudinal grooves. Dactylus of

claw with 10-13 (usually 12-13) teeth L. tredecimdentata Holthuis, 1941

4. Rostral plate with low median carina, flanked by longitudinal grooves, or with median

depression. Dactylus of claw with 7-9 teeth. Apex of uropodal endopod light

L. sulcata, new species

Rostral plate with distinct median carina, not flanked laterally by longitudinal grooves.

Dactylus of claw with 8-11 (usually 10-11) teeth (fewer in large females). Apex of

uropodal endopod dark L. maculata (Fabricius, 1793)

Lysiosquilla capensis Hansen, 1895

FIGURE 11

Lysiosquilla capensis Hansen, 1895:74.—Holthuis, 1967:15

[synonymy, including larvae].—Manning, 1969:5, fig. 1

[description and figure].

DIAGNOSIS.—Rostral plate (Figure lla) cordiform,

length and width subequal, median carina absent.

Antennal protopod (Figure lla) with sharp anterior projection above articulation of antennal peduncle. Antennal scale (Figure lla) slender, length

about 3 times greatest width, scale outlined by dark

pigment. Dactylus of raptorial claw (Figure lid)

with 15-17 teeth. Dorsal tooth of carpus of claw

not markedly deflexed mesioventrally. Ventral keel

of eighth thoracic somite (Figure 116) produced

into posteriorly directed spine. Sixth abdominal

somite and telson (Figure 11/) smooth or slightly

wrinkled dorsally. Basal segment of uropod with

strong spine ventrally at articulation of endopod

(Figure lie), apex of endopod dark.

REMARKS.—Although this species is not represented in the materials reported here, a diagnosis

has been included to complement the comparative

illustrations of this species given below (Figure 11).

A more detailed description with illustrations has

been given in an earlier paper (Manning, 1969).

During a visit to the Universitetets Zoologiske

Museum, Copenhagen, in 1971, I found that the

holotype of this species, a male, TL 59 mm, from

Port Elizabeth, South Africa, was deposited there,

rather than in the Musee Zoologique, Strasbourg,

as reported by Holthuis (1967).

Lysiosquilla capensis is the only member of the

genus in which the rostral plate is smooth dorsally,

being neither longitudinally sulcate nor carinate.

It also has more teeth on the claw, 15-17, than any

other representative of the genus. Finally, the ventral spine on the basal segment of the uropod is

better developed in this species than in any other

in the genus.

This is a relatively small species of Lysiosquilla.

It does not approach L. maculata in maximum size.

Males 59-104 mm long and other specimens 40-96

mm long have been recorded in the literature

(Manning, 1969).

DISTRIBUTION.—Apparently restricted to South

African waters, in relatively shallow water, from

shore to 48 fathoms (88 meters).

Lysiosquilla maculata (Fabricius, 1793)

FIGURES 1-3, 9

Squilla maculata Fabricius, 1793:511.

Lysiosquilla Miersii De Vis, 1883:321.

Lysiosquilla maculata.—Holthuis, 1967:18 [complete synonymy].—Manning, 1968:36, fig. 12 [comparison with L.

tredecimden tata].

MATERIAL.—Honolulu, Oahu, Hawaiian Islands; Albatross,

leg.; 12 Nov 1896: 1$, TL 118 mm (USNM 21489). Same;

Albatross, leg.; 1901: 1$ , TL 162 mm (USNM 64899).

Honolulu market; Albatross, leg.; 26 Mar 1902: 1 $, TL

220 mm (USNM 64901). Lagoon, Palmyra Island, Line

Islands; F. C. Sibley et al., leg.; 15 May 1966: 1$, TL

140 mm (USNM 120230). Apia, Upolu Islands, Samoa

Islands: D. S. Jordan, leg.; 1902: 1 <J, TL 260 mm (USNM

64904). Data same: 2$ , TL 240-285 mm (USNM 64905).

Samoa Islands; A. B. Steinberger, leg.: 1$ , TL 298 mm

(USNM 5148). Tahiti, Society Islands; Mission Ranson, leg.;

1952: 1$ , TL 198 mm (USNM 124719). Papara, Tahiti,

Society Islands; Albatross, leg.; 11 Nov 1899: 19 . TL 190

mm (USNM 69592). Raiatea, Society Islands; J. M. Clements, leg.; 15 Aug 1927: 2$, TL 255-285 mm; 1 $ , TL

267 mm (USNM 62613). Farepiti Point, Bora Bora, Society

Islands; purchased from fishermen; Smithsonian-Bredin Sta

50-57; 23 Apr 1957: 2 $, TL 253-317 mm; 2$ , both TL

ca. 300 mm (USNM 105674). Nuku Hiva, Marquesas Islands;

W. H. Jones, leg.; 1883: 1 $, TL 210 mm (USNM 6593).

Toau Atoll, Tuamotu Archipelago; G. Pinchot, leg.; 5 Oct
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 1.—l.ysiosquilla macula/a (Fabricius), male, TL 76 mm, Zanzibar: a, anterior part of

body; b, ventral keel of eighth thoracic somite, lateral view; c, uropod, ventral view. Setae

omitted.

1929: 1 $, TL 222 mm (USNM 63164). CclelHS, Indonesia;

M. C. Raven, leg.; 1 9 , TL 175 mm (USNM 69595). Moreton Bay, Queensland, Australia; holotypc of L. miersii De

Vis: 19 , TL ca. 225 mm (Queensland Museum). Dar es

Salaam, Tanzania; Cmdr. Smart, R. N., leg.: 1 9 , TL 120

mm (BMNH 1919.11.11.2). Zanzibar; J. D. Robinson, Peace

Memorial Museum: 1J , TL 76 mm (BMNH 1953.5.28.2-3).

Diego Garcia Atoll, Chagos Archipelago; 07°26'S, 72°26'E;

shore- to 1.2 m, dipnet; H. A. Fchlmaun Sta HA67-42;

12 Jul 1967: 1 9 , TL 180 mm (USNM 168530).

DIAGNOSIS.—Rostral plate (Figure 9a) usually

cordiform, occasionally sub triangular, width usually

greater than length, with short, raised median

carina anteriorly. Antennal protopod (Figure 9a)

lacking anterior projection. Antennal scale (Figure

9a) broad, length less than 3 times greatest width.

Scale not outlined by dark pigment but with irregular dark spot on surface. Dactylus of claw

(Figure 9d) usually with 10-11 teeth, fewer and

smaller in large females. Dorsal tooth of carpus of

claw not deflexed mesioventrally. Sixth abdominal

somite and telson (Figure 9/) smooth or slightly

wrinkled dorsally. Basal segment of uropod (Figure

9e) unarmed ventrally at articulation of endopod,

apex of endopod dark.

DESCRIPTION.—Eye (Figures la, 2a,b, 3a) large,

cornea bilobed, set slightly obliquely on stalk.

Ocular scales erect, triangular, apices inclined

anteriorly.

Antennular peduncle short, about half as long

as carapace. Dorsal processes of antennular somite

visible lateral to rostral plate as sharp, anteriorly

directed spines.

Antennal scale (Figures la, 2a, 3a) broad, curved,

length about 2.5 times greatest width. Scale with

large dark spot on surface. Antennal peduncle not

extending beyond eye. Antennal protopod lacking

triangular anterior projection, 1 mesial and 2 ventral papillae present.

Rostral plate usually cordiform (Figures la, 2a),

occasionally subtriangular (Figure 3a), width usually greater than length. Short median carina present on anterior half or third of plate, carina not

flanked laterally by deep longitudinal grooves; carina low, broad, less distinct, and with apex blunter

in large specimens.

Dactylus of raptorial claw with 8-11 teeth, usually 10-11; number and size of teeth reduced in

large females (TL 186-200 mm or larger); outer

margin of dactylus sinuate. Propodus of claw longer

than carapace except in large females in which it

may be shorter. Propodus and carpus comparatively

more inflated in large females and males; in large

females, complement of proximal movable teeth on

opposable margin of carpus reduced and carpus

and propodus ornamented with hairs. Dorsal tooth

of carpus not deflexed mesioventrally.

Mandibular palp and 5 epipods present.

Ventral keel of eighth thoracic somite rounded

(Figures 16, 2c).

Abdomen smooth, unarmed. Sixth somite smooth
NUMBER 259

FIGURE 2.—Lysiosquilla maculata (Fabricius), holotype of Lysiosquilla miersii De Vis, female,

TL ca. 225 mm, Queensland: a, anterior part of body; b, eye; c, ventral keel of eighth thoracic

somite, lateral view; d, uropod, ventral view. Setae omitted.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Lysiosquilla metadata (Fabricius), female, TL 180 mm, Diego Garcia Atoll: a,

anterior part of body; b, rostral plate. Setae omitted.

medially, with low, broad lateral boss flanked mesially by shallow longitudinal groove. Sixth somite

with triangular lobe ventrolaterally anterior to

articulation of each uropod.

Telson much broader than long, smooth or

slightly wrinkled, with broad, shallow pits dorsally,

with raised, smooth, subtriangular or ovate median

boss and lower submedian bosses on dorsal surface.

Lateral margin of telson lacking lateral carina in

large specimens, with low lateral carina on proximal fourth in smaller ones (male, TL 76 mm).

Posterior margin of telson with 3-4 pairs of fixed

projections, inner indistinct, outer sharpest.

Basal segment of uropod with slender dorsal

spine. Proximal segment of exopod shorter than

distal, with 8 movable spines, distalmost not exceeding midlength of distal segment. Endopod

broad, length about or less than twice greatest

width; apex of endopod dark. Basal prolongation

consisting of 2 spines, trefoil in cross section, inner

longer. Ventral surface of uropodal protopod unarmed at articulation of endopod (Figure 2d).

COLOR.—Eyestalks with scattered dark chromatophores. Anterior half of rostral plate with dark

spot. Anterior margin of antennal protopod and

dorsal surface of antennal peduncle with scattered

dark chromatophores; scale (Figures la, 3a) with

large, irregular dark spot on surface, margin not

outlined by dark pigment. Claw, when folded, with

3 broad bands of dark pigment, propodus without

separate distal vertical dark bar. Carapace with 3

broad, dark bands. Posterior 3 thoracic and all

abdominal somites with diffuse anterior and more

concentrated posterior dark bars, posterior bar on

margin. Sixth abdominal somite with dark lateral

patch extending onto basal segment of uropod.
NUMBER 259

Telson with median and 2 submedian dark spots

on posterior half, spots distinct or coalesced anteriorly. Basal segment of uropod dark proximally.

Uropodal exopod with dark spot overlapping segments, apex of distal segment light (Figure 1c).

Distal two-thirds of endopod dark (Figure lc).

MEASUREMENTS.—Males, TL 76-317 mm; females,

TL 120-300 + mm. Other measurements of a male,

TL 317 mm: carapace length 52.9; cornea width

11.4; antennal scale length 36.6, width 17.3; rap

torial propodus length 81.2; fifth abdominal somite

width 64.2. Other measurements of a female, TL

300 + mm: carapace length 56.2; cornea width 9.4;

antennal scale length 28.4, width 16.2; raptorial

propodus length 49.3; fifth abdominal somite width

71.3.

This is the largest known stomatopod. Females

as long as 385 mm have been recorded in the literature; specimens longer than 250 mm are not uncommon in either sex.

REMARKS.—Lysiosquilla maculata is the largest,

the most common, and the most widely distributed

species of the genus in the Indo-West-Pacific region.

It resembles L. sulcata, new species, described below, and differs from the other species in that region in having a broad antennal scale, a rounded

ventral keel on the eighth thoracic somite and in

lacking an angled dorsal projection on the antennal

protopod. It differs from L. sulcata in having more

teeth on the claw (usually 10-11 rather than 7-9),

a broader rostral plate lacking longitudinal grooves

along the median carina, and a dark apex on the

uropodal endopod.

In general, the rostral plate in L. maculata is

cordiform, with a short anterior carina that may

be very low in large specimens. In some specimens,

however, the plate is subtriangular (Figure 3a) or

even pentagonal, approaching that of L. sulcata in

shape. Variation in the shape of the plate is not

unusual in this species.

Through die kindness of Dr. Bruce Campbell,

Queensland Museum, I was able to examine die

holotype of Lysiosquilla miersii De Vis. As suggested by Stephenson (1953), it can be identified

with L. maculata. Sketches of the holotype are

shown in Figure 2.

Most of die references included in Holthuis'

(1967) complete synonymy for L. maculata require

verification, which can be accomplished only by

examining the specimens on which they are based.

Certainly the earlier records are not reliable and

may apply to any member of the genus occurring

in the Indo-West-Pacific region.

SEXUAL DIMORPHISM.—Several authors, including

Kemp (1913), Bigelow (1931), and Holthuis (1941),

have reported that large females of this species,

about 186 mm in total length or larger, exhibit

marked secondary sexual characters. In such specimens the eye size is reduced and the propodus of

the raptorial claw is comparatively smaller than in

smaller specimens. In addition, the propodus is

dilated, the proximal movable spines on the propodus are reduced and some may be absent, the

propodus and carpus may be ornamented with numerous hairs, and the teeth on the dactylus of the

claw are reduced in number as well as in size. The

reduction in number of teeth in large females of

L. maculata has led to some confusion in the past,

for such females may have the same number of

teeth on the claw as L. sulcirostris Kemp, 1913 (and

L. sulcata, new species). Lysiosquilla maculata var.

sulcirostris was described by Kemp from a male,

113.5 mm long, with a narrow, grooved rostral

plate and 8 teeth on die dactylus of die claw. The

similarity in number of teeth on the claw in large

females of L. maculata and in L. sulcirostris led

Monod (1925) and Bigelow (1931) to question die

validity of Kemp's variety and further induced

Chopra (1934) to publish a long discussion of these

characters based on specimens identified with L.

maculata in the collection of the Indian Museum.

Monod and Bigelow were basing their observations

on highly modified females of L. maculata.

The specimen from Mauritania that Monod

(1925) identified with L. sulcirostris represents a

new species, the description of which is in press

(Manning).

DISTRIBUTION.—Widely distributed in the IndoWest-Pacific region, from Japan to die western Indian Ocean, usually in shallow water. Most literature records require verification.

Lysiosquilla sulcata, new species

FIGURES 4-6, 10

Lysiosquilla sulcirostris.—Manning, 1970:1438, fig. S [not

Lysiosquilla sulcirostris Kemp, 1913].

MATERIAL.—Saipan, Marianna Islands; S. Hofer, leg.; 1918;

paratype: 1 $, TL 150 mm (SMF). Pacific Ocean; no other
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

data; paratype: 1 $, TL 92 mm (USNM 124770). Konori

Island, Mios Wendi, Padaido Group, West New Guinea

(West Irian); National Science Foundation sta 464; 28

Jan 1956; paratype: 1^, TL 122 mm (RMNH). Amboina,

Indonesia; Strubell, leg.; paratype: 1$, TL 75 mm (SMF).

Rasdu Atoll, near Weligandu Island (04°18'N, 73°OO'E),

Maldive Islands; sand zone with coral; Klauscwitz, leg.; 19

Mar 1958; paratype: 1 ?, TL 98 mm (SMF). Between

Manlefuri and Mara Islands, Fadiffolu Atoll, Maldive Islands;

05°21'SO"N, 73°25'15"E; J. S. Garth, R. Robertson, F. C.

Ziesenhenne, leg.; HOE, Sta GA 64-25; 23-24 Mar 1964;

holotype: 1$, TL 99 mm (USNM 156253). Grand Recif,

Tulear, Madagascar; between buoys; coarse sand, plants, in

shallow water; R. Derijard, leg.; 25 Apr 1967; paratype:

1 $, TL 41 mm (MNHNP). Zanzibar; no other data; paratype: 1 $, TL 129 mm (BMNH 1935.558.2-3).

DIAGNOSIS.—Rostral plate (Figure 10a) subtriangular or subpentagonal, length and width subequal or length slightly greater, with median longitudinal depression anteriorly, median carina

reduced or absent. Antennal protopod (Figure 10a)

lacking anterior projection. Antennal scale (Figure

10a) broad, length less than 3 times greatest width.

Scale not outlined by dark pigment but with scattered dark spots on surface. Dactylus of raptorial

claw (Figure lOd) with 7-8 teeth, dorsal tooth of

carpus of claw not deflexed mesioventrally. Ventral

keel of eighth thoracic somite (Figure 10&) rounded.

Sixth abdominal somite and telson (Figure 10/)

smooth or with shallow pits dorsally. Basal segment

of uropod (Figure 10c) unarmed ventrally at articulation of endopod, apex of endopod light.

DESCRIPTION.—Eye (Figures 4a,b, 5a, 6) large,

cornea bilobed, set slightly obliquely on stalk.

Ocular scales erect, triangular, apices inclined anteriorly.

Antennular peduncle short, about half as long

as carapace. Dorsal processes of antennular somite

visible lateral to rostral plate as broad, anteriorly

directed spines.

FIGURE 4.—Lysiosquilla sulcata, new species, male holotype, TL 99 mm, Maldive Islands: a,

anterior part of body; b, eye; c, rostral plate; d, ventral keel of eighth thoracic somite, lateral

view; e, uropod, ventral view. Setae omitted.
NUMBER 259

FIGURE 5.—Lysiosquilla sulcata, new species, male paratype, TL 129 mm, Zanzibar: a, anterior

part of body; b, rostral plate; c, ventral keel of eighth thoracic somite, lateral view; d, uropod,

ventral view (outer spine of basal prolongation broken). Setae omitted.

Antennal scale broad (Figures 4a, 5a, 6), length

about 2.5 times greatest width. Scale not outlined

by dark pigment, with dark spots on surface. Antennal peduncle not extending beyond eye. Antennal protopod (Figures 4a, 5a, 6) lacking anterior

projection, with 1 mesial and 2 ventral papillae.

Rostral plate (Figures 4a, 5a, 6) subtriangular or

subpentagonal, length and width subequal or

length greater, apex deflexed. Plate longitudinally

sulcate anteriorly, occasionally with obscure, low

carina in channel (Figures 4a, 6).

Dactylus of raptorial claw with 7-9 teeth, outer

margin of dactylus sinuate. Teeth of claw not

markedly reduced in large females available for

study. Propodus of claw longer than carapace in

males, slightly shorter than carapace and ornamented with simple hairs in female. Dorsal tooth

of carpus of claw not deflexed mesioventrally.

Mandibular palp and 5 epipods present.

Ventral keel of eighth thoracic somite rounded

(Figures 4d, 5c).

Abdomen smooth, unarmed. Sixth somite with

smooth lateral boss on each side, flanked mesially

by shallow, longitudinal groove. Sixth somite with

blunt triangular projection ventrolaterally anterior

to articulation of each uropod.

Telson broader than long, pitted dorsally, with

low, triangular, smooth median boss and 2 lower

submedian bosses on dorsal surface. Lateral margin

of telson not distinctly carinate. Posterior margin

of telson with 3 pairs of fixed, blunt projections,

outer sharpest.

Basal segment of uropod with dorsal spine. Proximal segment of exopod shorter than distal, with

8 slender movable spines, distalmost extending to

midlength of distal segment. Endopod broad,

length about twice greatest width. Apex of endopod

light in color (Figures 4e, 5d). Basal prolongation
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of uropod consisting of 2 spines, trefoil in cross

section. Ventral surface of basal segment of uropod

unarmed at articulation of endopod (Figures 4e,

5d).

COLOR.—Largely faded on all specimens. Rostral

plate and antennal scale each with diffuse dark

spot, appearing speckled. Carapace with 3 dark

bands. Posterior 3 thoracic and all abdominal somites with posterior margin outlined by dark pigment. Telson with 3 dark spots dorsally. Uropod

with dark spot dorsally on basal segment and dark

spot on exopod at articulation of distal segment;

endopod with dark spot on outer surface, inner

margin and apex light (Figures 4e, 5d).

MEASUREMENTS.—Males, TL 41-150 mm; females,

TL 75-98 mm. Other measurements of male holotype, TL 99 mm: carapace length 17.3; cornea width

5.7; antennal scale length 12.7, width 5.3; raptorial

propodus length 24.1; fifth abdominal somite with

18.7.

REMARKS.—Lysiosquilla sulcata, new species, resembles L. maculata and differs from the other

Indo-West-Pacific species of the genus in lacking

an anterior projection on the antennal protopod,

and in having a rounded ventral keel on the eighth

thoracic somite as well as a broad rather than

slender antennal scale. It differs from L. maculata

in having fewer teeth on the claw, a light apex on

the uropodal endopod, and in the structure of the

rostral plate. In L. sulcata the plate is either longitudinally sulcate or bears a short anterior carina

flanked laterally by longitudinal grooves. The rostral plate of L. sulcata resembles that of L. sulcirostris (see below), but Kemp's species has a slender

antennal scale, a projection on the antennal protopod, an angled ventral keel on the eighth thoracic somite, and a dark apex on the uropodal

endopod.

The shape of the rostral plate in the small male

from Tulear, the specimen which I (Manning,

1970) had identified with L. sulcirostris, is quite

different from that illustrated here and is reproduced for comparison (Figure 6). Apparently in

this species, as in L. maculata, the shape of the

plate is variable. In some specimens the anterior

carina is so poorly developed that the plate appears

to have a single median, longitudinal groove.

ETYMOLOGY.—The name is from sulcatus (Latin,

= grooved), alluding to the structure of the rostral

plate.

DISTRIBUTION.—Indo-West-Pacific region, from

scattered localities between Saipan and Zanzibar.

It apparently lives in shallow water.

FIGURE 6.—Lysiosquilla sulcata, new species, male paratype, TL 41 mm, Tulear: anterior part

of body. Setae omitted.
NUMBER 259 11

Lysiosquilla sulcirostris Kemp, 1913

FIGURES 7-8, 12

Squilla maculata.—De Haan, 1849:221 [not Squilla maculata

Fabricius, 1793].

Lysiosquilla maculata var. sulcirostris Kemp, 1913:116, pi.

8: figs. 92-93.—Holthuis, 1941:272.—Serene, 1954:6, 7,

8, 11, IS, 70, figs. 1, 3, pi. 5: figs. 3-4, pi. 6: figs. 3-4.

Lysiosquilla sulcirostris.—Holthuis, 1967:23 [complete synonymy].

MATERIAL.—Japan; P. F. Von Siebold, leg.: 1$, TL 58

mm (RMNH 12S). Insel Lombussa (Jesus-Maria), Admiralty

Group; H. Schroede, leg.; 3 Mar 1910: 1 $, TL 325 mm

(ex ZMB 13774). Lembeh Strait, N. Celebes, Indonesia;

Oct 1941: 1 postlarva, TL 20 mm (RMNH 264S). Celebes,

Indonesia; 29 Jan 1894: 1 $, TL 134 mm (NMV)., Batavia

(Djakarta), Indonesia; Bleeker, leg.; 1 $, TL 105 mm

(MNHNP). Andaman Islands; Homfray and Tytler, don.;

holotype: 1$, TL 113.5 mm (ZSI 7564/10). Tany Kely,

NW coast of Madagascar (Malagasy Republic); 20 m; P.

Laboute, leg.; 14 Jan 1971: 1£ , TL 115 mm (USNM

168531).

DIAGNOSIS.—Rostral plate (Figure 12a) subtriangular, appearing elongate, usually slightly broader

than long, median carina on anterior part flanked

laterally by deep longitudinal groove. Antennal

protopod (Figure 12a) with blunt triangular anterior projection above articulation of antennal peduncle. Antennal scale (Figure 12a) slender, length

about 3 times greatest width. Scale covered with

dark, pigment. Dactylus of raptorial claw (Figure

12d) with 7-8 teeth. Dorsal tooth of carpus of claw

not deflexed mesioventrally. Ventral keel of eighth

thoracic somite (Figure 126) produced into angled

lobe or sharp, posteriorly directed spine. Sixth abdominal somite and telson (Figure 12/) smooth or

slightly pitted dorsally. Basal segment of uropod

unarmed ventrally at articulation of endopod (Figure 12e), apex of endopod dark.

DESCRIPTION.—Eye (Figures 7, Sa,b) large,

cornea bilobed, set slightly obliquely on stalk. Ocular scales erect, sharp, apices inclined anteriorly.

Antennular peduncle short, slightly more than

half as long as carapace. Dorsal processes of antennular somite visible lateral to rostral plate as sharp,

anterolaterally directed spines.

FIGURE 7.—Lysiosquilla sulcirostris Kemp, male holotype, Andaman Islands, TL 113.5 mm:

a, anterior part of body; b, eye. Setae omitted.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Antennal scale (Figures la, 8a) slender, curved,

length more than 3 times greatest width; scale with

dark pigment scattered over surface. Antennal protopod with triangular projection dorsally (Figures

la, 8a) and with 1 mesial and 2 ventral papillae.

Rostral plate (Figures la, %a,c) triangular, appearing elongate but usually slightly broader than

long; median carina on anterior half flanked on

each side by deep longitudinal groove.

Dactylus of claw with 7-8 teeth, outer margin

of dactylus straight or slightly sinuate. Propodus

of claw longer than carapace [except perhaps in

adult females (Serene, 1954)]. Dorsal tooth of carpus not deflexed mesioventrally.

Mandibular palp and 5 epipods present.

Ventral keel of eighth thoracic somite produced

posteriorly into angled lobe or sharp spine (Figure 8d).

Abdomen smooth, unarmed. Sixth somite with

broad lateral boss flanked mesially by shallow

groove and with triangular lobe ventrolaterally

anterior to articulation of each uropod.

Telson much broader than long, slightly pitted

dorsally, with low, smooth, triangular median boss

and 2 lower submedian bosses on dorsal surface.

Lateral margin of telson not distinctly carinate.

Posterior margin of telson with 4 pairs of fixed

projections, outer 2 sharpest.

Basal segment of uropod with dorsal spine. Proximal segment of exopod shorter than distal, with

7-8 slender, movable spines, distalmost extending

about to midlength of distal segment. Endopod

broad, apex dark (Figure 8e). Basal prolongation

consisting of 2 slender spines, trefoil in cross section, inner longer. Ventral surface of uropodal protopod unarmed at articulation of endopod ( Figure Se).

COLOR.—Antennal scale with scattered dark chromatophores on surface, not forming dark spot. Carapace with 3 dark bands. Posterior 3 thoracic and

anterior 5 abdominal somites with anterior and

posterior dark bars, posterior margins outlined in

black. Telson with dark bar across midlength, darkest laterally. Basal segment of uropod with dark

anterior patch. Dark spot at articulation of segments of uropodal exopod covering proximal portion of distal segment. All but proximal fifth or

less of uropodal endopod dark (Figure 8e).

MEASUREMENTS.—Males, TL 58-325 mm; no females examined; postlarva, TL 20 mm. Other

measurements of male holotype, TL 113.5 mm:

carapace length 19.7; cornea width 6.0; antennal

FIGURE 8.—Lysiosquilla sulcirostris Kemp, male, TL 58 mm, Japan: a, anterior part of body;

b, eye; c, rostral plate; d, ventral keel of eighth thoracic somite, lateral view; e, uropod,

ventral view. Scute omitted.
NUMBER 259 13

scale length 11.0, width 3.6; raptorial propodus

length 28.1; fifth abdominal somite width 22.6.

REMARKS.—Lysiosquilla sulcirostris resembles L.

capensis Hansen and L. tredecimdentata Holthuis

and differs from L. maculata (Fabricius) and L. sulcata, new species, in having an anterior projection

on the antennal protopod, a slender antennal scale,

and a posteriorly sharp ventral projection on the

eighth thoracic somite. It differs from L. capensis

and L. tredecimdentata in having fewer teeth on

the claw (7-8 rather than 15-17 or 10-13) and a

rostral plate in which the carina is flanked laterally

by deep longitudinal grooves.

Lysiosquilla sulcirostris also resembles an undescribed West African species (Manning, in press)

that Monod (1925) reported as L. maculata var.

sulcirostris. The West African species differs in

having a spine on the ventral surface of the uropodal protopod at the articulation of the endopod.

In other respects they are very similar.

The largest male recorded here, total length 325

mm, was in a lot with several specimens of L.

maculata of similar size; it is the largest known

specimen of the species.

Marked sexual dimorphism, similar to that observed for L. maculata, has been recorded for this

species by Serene (1954). In a female 143 mm long,

Serene noted a reduction in size of the eye and the

number of teeth on the claw (5 rather than 8), an

enlargement of the propodus of the claw, and the

presence of tufts of hairs on the propodus and

carpus. He pointed out that in this species eye size

increased until specimens had attained a length of

70 to 90 mm; in larger specimens, the eye size of

females decreased with increasing total length. No

reduction in length of the propodus of the claw

was reported by Serene. These data indicate that

L. sulcirostris matures at a much smaller size than

does L. maculata.

DISTRIBUTION.—Indo-West-Pacific region, from

scattered localities between Japan and Madagascar. All previous records (see Holthuis, 1967) require verification.

Lysiosquilla tredecimdentata Holthuis, 1941

FIGURE 13

Lysiosquilla maculata.—Kemp, 1913:111, pi. 8: figs. 87-89

[part].—Chopra, 1934:28 [part ?; material not listed] [not

L. maculata (Fabricius, 1793)].

Lysiosquilla maculata var. tredecimdentata Holthuis, 1941:

273, fig. 6.

Lysiosquilla tredecimdentata.—Holthuis, 1967:23 [synonymy].

—Manning, 1968:38, fig. 13 [description]; 1969:6.

^Lysiosquilla maculata.—Alikunhi, 1967:904. ligs. 158-167

[postlarval development] [not L. maculata (Fabricius,

1793)].

MATERIAL.—Coconada, South India; G. W. Wicks, don.:

5 3 , TL 72-1325 mm; 3$, TL 106-186 mm (/SI 3517-19/7

and 3528-31/7). Madras, India; 2$, TL 138-144 mm (ZSI

3630-31/7). Andaman Islands: 1 fragmented $ (ZSI 3044/5).

No data: 1 $, TL 91 mm (ZSI 3037/5).

AH of these specimens were identified as L. maculata by

Kemp (1913).

DIAGNOSIS.—Rostral plate (Figure 13a) cordiform,

broader than long, with median carina on anterior

third. Antennal protopod (Figure 13a) with triangular anterior projection above articulation of antennal peduncle. Antennal scale (Figure 13a) slender, length at least 3 times greatest width. Scale

outlined by dark pigment. Dactylus of raptorial

claw (Figure 13d) with 10-13, usually 12-13, teeth.

Dorsal tooth of carpus of claw deflexed mesioventrally. Ventral keel of eighth thoracic somite (Figure 136) produced into posteriorly directed spine.

Sixth abdominal somite and telson (Figure 13/)

smooth or slightly wrinkled dorsally. Basal segment

of uropod occasionally armed with small spinule

ventrally at articulation of endopod (unarmed in

Figure l$e), apex of endopod dark.

REMARKS.—Lysiosquilla tredecimdentata is a distinctive species that resembles L. capensis and L.

sulcirostris and differs from L. maculata and L.

sulcata, new species, in having a slender antennal

scale, outlined in dark pigment, an anterior projection on the antennal protopod, and a posterior

spine on the ventral keel of the eighth thoracic

somite. It differs from L. capensis in having a

broader rostral plate with a distinct dorsal carina

and 10-13 rather than 15-17 teeth on the claw.

Lysiosquilla tredecimdentata differs from L. sulcirostris in having a broader rostral plate that lacks

deep grooves flanking the median carina and in

having 10-13 rather than 7-8 teeth on the dactylus of the claw.

This is a relatively large species. Males ranging

in length from 120 to 259 mm and females 116 to

246 mm long have been recorded in the literature.

In the Indo-West-Pacific region, only L. maculata

and L. sulcirostris, both of which may exceed 300

mm in total length, may attain a larger size.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 9-13.—Indo-West-Pacific species of Lysiosquilla: 9, L. maculata (Fabricius); 10, L.

sulcata, new species; 11, L. capensis Hansen; 12, L. sukirostris Kemp; 13, L. tredecimdentata

Holthuis. (Figures 9-13: a, anterior part of body; b, ventral keel of eighth thoracic somite,

lateral view; c, eye; d, claw; e, uropod, ventral view; /, sixth abdominal somite, telson, and

uropod.)
NUMBER 259 15

Alikhunhi (1967) published a detailed account of

the postlarval development of two species of Lysiosquilla from Madras, India. These he identified as

L. maculata and L. sulcirostris. As I pointed out

earlier (Manning, 1970), the features listed by Alikunhi as characteristic of his young specimens of

L. maculata (10-12 teeth on the claw, a slender

antennal scale outlined by dark pigment) actually

are diagnostic for L. tredecimdentata, raising the

possibility that he was dealing with specimens of

both L. maculata and L. tredecimdentata or with

the latter species only. His material should be

reexamined.

All of the material reported here was identified

by Kemp (1913) as L. maculata. If his illustrations

(pi. 8: figs. 87-89) are accurate, the specimens illustrated also are referable to L. tredecimdentata

rather than to L. maculata; the broad dark bands

on the body and the slender antennal scales are

characteristic of the former species.

DISTRIBUTION.—Now known with certainty from

localities in the western Indian Ocean, from the

Red Sea southward to South Africa, eastward to

southern India and the Andaman Islands. It may

be expected to have a much wider distribution

than now recognized.
Literature Cited

Alikunhi, K. H.

1967. An Account of the Post-larval Development, Moulting, and Growth of the Common Stomatopods of

the Madras Coast. Pages 824-939 in Part II in

Proceedings of the Symposium on Crustacea, Marine Biological Association of India, figures 1-194,

plates 1-3.

Bigelow, R. P.

1931. Stomatopoda of the Southern and Eastern Pacific

Ocean and the Hawaiian Islands. Bulletin of the

Museum of Comparative Zoology, Harvard University, 72(4): 105-191. figures 1-10, plates 1-2.

Chopra, B.

1934. On the Stomatopod Crustacea Collected by the

Bengal Pilot Service off the Mouth of the River

Hughli, Together with Notes on Some Other Forms.

Records of the Indian Museum, 36:17-43, figures

1-5.

De Vis, C. W.

1883. Description of a Species of Squill from Moreton

Bay. Proceedings of the Linnean Society of New

South Wales, 7:321-322.

Fabricius, J. C.

1793. Entomologia systematica emendata et aucta, secundum classes, ordines, genera, species, adjectis synonimis, locis, observationibus, descriptionibus. Volume 2, pages ir-viii + 1-519.

Haan, W. De

1833-1850. Crustacea. Pages i-xvii, i-xxxi, ix-xvi, 1-243 in

Von Siebold, Fauna Japonica sive descriptio Animalium, quae in itinere per Japoniam, jussu et Auspiciis

Superiorum, qui Summum in India Batava Imperium Tenent, suscepto, annis 1823-1930 collegit,

notis observationibus, et adumbrationibus illustravit. Plates A-J, L-Q, 1-55, circ. tab. 2- LugduniBatavorum: A. Arnz.

Hansen, H. J.

1895. Isopoden, Cumaceen und Stomatopoden der

Plankton-Expedition. Pages 1-105 in volume 2

(Gc) in Ergebnisse der Plankton-Expedition der

Humboldt-Stiftung. Plates 1-8.

Hobbs, Horton H., Jr.,

1974. A Checklist of the North and Middle American

Crayfishes (Decapoda: Astacidae and Cambaridae).

Smithsonian Contributions to Zoology, 166:1-161,

figures 1-294.

Holthuis, L. B.

1941. The Stomatopoda of the Snellius Expedition. In

Biological Results of the Snellius Expedition, XII.

Temminckia, 6:241-294, figures 1-9.

1967. Fam. Lysiosquillidae et Bathysquillidae: Stomatopoda I. Pages 1-28 in volume 1 in Gruner and

Holthuis, editors, Crustaceorum Catalogus. The

Hague: W. Junk.

Kemp, S.

1913. An Account of the Crustacea Stomatopoda of the

Indo-Pacific Region, Based on the Collection in

the Indian Museum. Memoirs of the Indian

Museum, 4(1): 1-217, 10 figures, plates 1-10.

Manning, Raymond, B.

1968. Stomatopod Crustacea from Madagascar. Proceedings of the United States National Museum, 124

(3641):1-61, figures 1-16.

1969. Notes on Some Stomatopod Crustacea from Southern Africa. Smithsonian Contributions to Zoology,

1:1-17, figures 1-4.

1970. Some Stomatopod Crustaceans from Tulear, Madagascar. Bulletin du Musium National d'Histohe

Naturelle, (2)41(6)(1969):1429-1441, figures 1-3.

In press. A Monograph of the West African Stomatopod

Crustaceans. Atlantide Reports (Copenhagen).

Michel, A.

1970. Larves Pelagiques et Post-Larves du Genre Lysiosquilla (Crustaces Stomatopodes) dans le Pacifique

Tropical Sud et Equatorial. Cahiers ORSTOM,

Serie Oceanographique, France, 8(3):53-75, figures

1-10.

Monod, Th.

1925. Sur les Stomatopodes de la Cote Occidentale

d'Afrique. Bulletin de la Societi des Sciences Naturelles du Maroc, 5(3):86-93, plates 20-21.

Serene, R.

1954. Observations Biologiques sur les Stomatopodes.

Mimoires de I'Institut Oceanographique de Nhatrang, 8:1-93, figures 1-15, plates 1-10.

Stephenson, W.

1953. Notes on the Australian Stomatopoda (Crustacea)

in the Collections of the Queensland Museum.

Memoirs of the Queensland Museum, 13(l):40-49.

16
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