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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 259

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. maculata (Fabricius) and L. sulcirostris 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 PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnacus).

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′08s [595′.382] 77–10934

Synopses of the Indo-West-Pacific Species of Lysiosquilla Dana, 1852 (Crustacea: Stomatopoda: Lysiosquillidae)

Raymond B. Manning

____________

Raymond B. Manning, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560.

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 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 11a, 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 11a, 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-West-Pacific species and may well occur in all species. Kemp (1913) and Bigelow (1931), among others, have reported on dimorphism in L. maculata, and Serène (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-West-Pacific 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

Muséum National d'Histoire Naturelle, Paris

NMV

Naturhistorisches Museum, Vienna

RMNH

Rijksmuseum van Natuurlijke Historie, 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 Humboldt-Universität 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, Muséum 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 Historie, 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. Türkay 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

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 11a) cordiform, length and width subequal, median carina absent. Antennal protopod (Figure 11a) with sharp anterior projection above articulation of antennal peduncle. Antennal scale (Figure 11a) slender, length about 3 times greatest width, scale outlined by dark pigment. Dactylus of raptorial claw (Figure 11b) with 15–17 teeth. Dorsal tooth of carpus of claw not markedly deflexed mesioventrally. Ventral keel of eighth thoracic somite (Figure 11b) produced into posteriorly directed spine. Sixth abdominal somite and telson (Figure 11f) smooth or slightly wrinkled dorsally. Basal segment of uropod with strong spine ventrally at articulation of endopod (Figure 11e), 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 Musée 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. tredecimdentata].

MATERIAL.—Honolulu, Oahu, Hawaiian Islands; Albatross, leg.; 12 Nov 1896: 1 art, TL 118 mm (USNM 21489). Same; Albatross, leg.; 1901: 1 art, TL 162 mm (USNM 64899). Honolulu market; Albatross, leg.; 26 Mar 1902: 1 art, TL 220 mm (USNM 64901). Lagoon, Palmyra Island, Line Islands; F. C. Sibley et al., leg.; 15 May 1966: 1 art, TL 140 mm (USNM 120230). Apia, Upolu Islands, Samoa Islands: D. S. Jordan, leg.; 1902: 1 art, TL 260 mm (USNM 64904). Data same: 2 art, TL 240–285 mm (USNM 64905). Samoa Islands; A. B. Steinberger, leg.: 1 art, TL 298 mm (USNM 5148). Tahiti, Society Islands; Mission Ranson, leg.; 1952: 1 art, TL 198 mm (USNM 124719). Papara, Tahiti, Society Islands; Albatross, leg.; 11 Nov 1899: 1 art, TL 190 mm (USNM 69592). Raiatea, Society Islands; J. M. Clements, leg.; 15 Aug 1927: 2 art, TL 255–285 mm; 1 art, TL 267 mm (USNM 62613). Farepiti Point, Bora Bora, Society Islands; purchased from fishermen; Smithsonian-Bredin Sta 50–57; 23 Apr 1957: 2 art, TL 253–317 mm; 2 art, both TL ca. 300 mm (USNM 105674). Nuku Hiva, Marquesas Islands; W. H. Jones, leg.; 1883: 1 art, TL 210 mm (USNM 6593). Toau Atoll, Tuamotu Archipelago; G. Pinchot, leg.; 5 Oct 1929: 1 art, TL 222 mm (USNM 63164). Celebes, Indonesia; M. C. Raven, leg.; 1 art. TL 175 mm (USNM 69595). Moreton Bay, Queensland, Australia; holotype of L. miersii De Vis: 1 female;, TL ca. 225 mm (Queensland Museum). Dar es Salaam, Tanzania; Cmdr. Smart, R. N., leg.: 1 art, TL 120 mm (BMNH 1919.11.11.2). Zanzibar; J. D. Robinson, Peace Memorial Museum: 1 art, 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. Fehlmann Sta HA67–42; 12 Jul 1967: 1 art, TL 180 mm (USNM 168530).

FIGURE 1.—Lysiosquilla maculata (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.

DIAGNOSIS.—Rostral plate (Figure 9a) usually cordiform, occasionally subtriangular, 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 9f) smooth or slightly wrinkled dorsally. Basal segment of uropod (Figure 9e) unarmed ventrally at articulation of endopod, apex of endopod dark.

DESCRIPTION.—Eye (Figures 1a, 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 1a, 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 1a, 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 1b, 2c).

Abdomen smooth, unarmed. Sixth somite smooth medially, with low, broad lateral boss flanked mesially by shallow longitudinal groove. Sixth somite with triangular lobe ventrolaterally anterior to articulation of each uropod.

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.

FIGURE 3.—Lysiosquilla maculata (Fabricius), female, TL 180 mm, Diego Garcia Atoll: a, anterior part of body; b, rostral plate. Setae omitted.

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 1a, 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. 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 1c).

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; raptorial 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 the kindness of Dr. Bruce Campbell, Queensland Museum, I was able to examine the 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 the 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 the dactylus of the 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 the 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 Indo-West-Pacific region, from Japan to the 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. 3 [not Lysiosquilla sulcirostris Kemp, 1913].

MATERIAL.—Saipan, Marianna Islands; S. Hofer, leg.; 1918; paratype: 1 art, TL 150 mm (SMF). Pacific Ocean; no other data; paratype: 1 art, 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 art, TL 122 mm (RMNH). Amboina, Indonesia; Strubell, leg.; paratype: 1 art TL 75 mm (SMF). Rasdu Atoll, near Weligandu Island (04°18′N, 73°00′E), Maldive Islands; sand zone with coral; Klausewitz, leg.; 19 Mar 1958; paratype: 1 art, TL 98 mm (SMF). Between Mafilefuri and Mara Islands, Fadiffolu Atoll, Maldive Islands; 05°21′30″N, 73°25′15″E; J. S. Garth, R. Robertson, F. C. Ziesenhenne, leg.; IIOE, Sta GA 64–25; 23–24 Mar 1964; holotype: 1 art, 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 art TL 41 mm (MNHNP). Zanzibar; no other data; paratype: 1 art, TL 129 mm (BMNH 1935.5.28.2–3).

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 10d) with 7–8 teeth, dorsal tooth of carpus of claw not deflexed mesioventrally. Ventral keel of eighth thoracic somite (Figure 10b) rounded. Sixth abdominal somite and telson (Figure 10f) smooth or with shallow pits dorsally. Basal segment of uropod (Figure 10e) 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.

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 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.

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, pl. 8: figs. 92–93.—Holthuis, 1941:272.—Serène, 1954:6, 7, 8, 11, 13, 70, figs. 1, 3, pl. 5: figs. 3–4, pl. 6: figs. 3–4.

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

MATERIAL.—Japan; P. F. Von Siebold, leg.: 1 art, TL 58 mm (RMNH 12S). Insel Lombussa (Jesus-Maria), Admiralty Group; H. Schroede, leg.; 3 Mar 1910: 1 art, 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 art, TL 134 mm (NMV)., Batavia (Djakarta), Indonesia; Bleeker, leg.; 1 art, TL 105 mm (MNHNP). Andaman Islands; Homfray and Tytler, don.; holotype: 1 art, TL 113.5 mm (ZSI 7564/10). Tany Kely, NW coast of Madagascar (Malagasy Republic); 20 m; P. Laboute, leg.; 14 Jan 1971: 1 art, 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 12b) produced into angled lobe or sharp, posteriorly directed spine. Sixth abdominal somite and telson (Figure 12f) 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, 8a,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.

Antennal scale (Figures 7a, 8a) slender, curved, length more than 3 times greatest width; scale with dark pigment scattered over surface. Antennal protopod with triangular projection dorsally (Figures 7a, 8a) and with 1 mesial and 2 ventral papillae.

Rostral plate (Figures 7a, 8a,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 (Serène, 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 8e).

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 scale length 11.0, width 3.6; raptorial propodus length 28.1; fifth abdominal somite width 22.6.

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. Setate omitted.

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 Serène (1954). In a female 143 mm long, Serène 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 Serène. 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, pl. 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, figs. 158–167 [postlarval development] [not L. maculata (Fabricius, 1793)].

MATERIAL.—Coconada, South India; G. W. Wicks, don.: 5 art, TL 72–132.5 mm; 3 art, TL 106–186 mm (ZSI 3517–19/7 and 3528–31/7). Madras, India; 2 art, TL 138–144 mm (ZSI 3630–31/7). Andaman Islands: 1 fragmented art (ZSI 3044/5). No data: 1 art, TL 91 mm (ZSI 3037/5).

All 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 13b) produced into posteriorly directed spine. Sixth abdominal somite and telson (Figure 13f) smooth or slightly wrinkled dorsally. Basal segment of uropod occasionally armed with small spinule ventrally at articulation of endopod (unarmed in Figure 13e), 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.

FIGURES 9–13.—Indo-West-Pacific species of Lysiosquilla: 9, L. maculata (Fabricius); 10, L. sulcata, new species; 11, L. capensis Hansen; 12, L. sulcirostris 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; f, sixth abdominal somite, telson, and uropod.)

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 (pl. 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 i–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. Lugduni-Batavorum: 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 I 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 Tuléar, Madagascar. Bulletin du Muséum National d'Histoire 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 Pélagiques et Post-Larves du Genre Lysiosquilla (Crustacés Stomatopodes) dans le Pacifique Tropical Sud et Équatorial. Cahiers ORSTOM, Série Océanographique, France, 8(3):53–75, figures 1–10.

Monod, Th.

1925.   Sur les Stomatopodes de la Cote Occidentale d'Afrique. Bulletin de la Societé des Sciences Naturelles du Maroc, 5(3):86–93, plates 20–21.

Serène, R.

1954.   Observations Biologiques sur les Stomatopodes. Mémoires de l'Institut Océanographique 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(1):40–49.

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