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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 258

Notes on Some Species of the Falcatus Group of Gonodactylus (Crustacea: Stomatopoda: Gonodactylidae)

Raymond B. Manning

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978

ABSTRACT

Manning, Raymond B. Notes on Some Species of the Falcatus Group of Gonodactylus (Crustacea: Stomatopoda: Gonodactylidae). Smithsonian Contributions to Zoology, number 258, 15 pages, 13 figures, 1978.—Five species of the Falcatus group of Gonodactylus are described and illustrated. Gonodactylus glabrous Brooks, G. mutatus Lanchester, and G. ternatensis De Man are removed from the synonymy of G. falcatus (Forskål) and are redescribed from type-specimens; G. falcatus is redescribed from topotypical material; and G. randalli, new species, is described. Gonodactylus rotundus Borradaile is considered a synonym of G. mutatus Lanchester.

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 (Linnaeus).

Library of Congress Cataloging in Publication Data

Manning, Raymond B.

Notes on some species of the Falcatus group of Gonodactylus (Crustacea, Stomatopoda, Gonodactylidae).

(Smithsonian contributions to zoology; no. 258)

Bibliography: p.

1. Gonodactylus. 2. Crustacea—Classification. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology; no. 258.

QL1.S54 no. 258 [QL444.M375] 591′.08s [595′.382] 77–10754

Notes on Some Species of the Falcatus Group of Gonodactylus (Crustacea: Stomatopoda: Gonodactylidae)

Raymond B. Manning

____________

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

Introduction

In an earlier paper (Manning, 1968:42), I pointed out that the Indo-West-Pacific species of Gonodactylus comprise three groups of species: the Chiragra group, containing G. chiragra (Fabricius, 1781) and its allies, large species with broad ocular scales and three mid-dorsal carinae on the telson; the Demanii group, including G. demanii Henderson, 1893 and its allies, small species with narrow ocular scales and three mid-dorsal carinae on the telson; and the Falcatus group, containing G. falcatus (Forskål, 1775) and related species, moderate to large species with narrow ocular scales and five mid-dorsal carinae on the telson. The American species form the fourth group, the Oerstedii group, containing species related to G. oerstedii Hansen, 1895, all of which have an accessory carina on the mesial margin of the intermediate carina of the telson.

The Demanii group may well include representatives of two distinct groups, one with G. demanii and allies, containing species that often have modified telsons and uropods, and one with G. affinis De Man and related species, in which the telson closely resembles that found in the Chiragra group but in which the size at maturity is reduced.

Only two species currently are recognized in the Falcatus group: G. falcatus sensu lato, reported to have a wide distribution from the Red Sea and South Africa to Hawaii and Japan, and G. graphurus Miers, 1875, known only from Australasian waters. Until recently, G. falcatus had been known as G. glabrous Brooks, although the epithets glaber (Henderson, 1893 and others), graphurus (Müller, 1887 and others), and glabrus (Roxas and Estampador, 1930) also had been used for this species. Holthuis (1941) pointed out that the oldest available name for this species was G. falcatus (Forskål, 1775), a name generally overlooked or mistakenly thought to be a synonym of the distantly related G. chiragra (Fabricius).

Both Brooks (1886) and Kemp (1913) illustrated the major difference between G. falcatus (as G. glabrous) and G. graphurus, i.e., the transversely grooved abdominal somites in the latter species. In general, since 1913 the specific epithet graphurus has been applied to the grooved species that Ingle (1971) correctly attributed to Miers (1875) and that now is known to be restricted to Australasian waters (Manning, 1966). Inasmuch as this species is relatively well known and described (Manning, 1966; references to 1965), it is not treated in detail here. The grooved abdominal somites are shown below in Figure 2b.

The identity of G. falcatus is less clear, for it now is certain that several species are being confused under that name. The following names are available for species in the Falcatus group other than G. graphurus: G. falcatus (Forskål, 1775); G. glabrous Brooks, 1886; G. glabrous var. ternatensis De Man, 1902; G. chiragra var. mutatus Lanchester, 1903; G. glaber var. rotundus Borradaile, 1907; and G. bossorotundus Roxas and Estampador, 1930.

With the exception of G. falcatus, all of those names published before 1913 were synonymized with G. glabrous by Kemp (1913), and virtually all subsequent authors followed Kemp in recognizing a single taxon that Holthuis (1941) called G. falcatus. One exception to this was Serène (1954), who recognized two taxa from Viet Nam, G. falcatus and a dwarf variety (“race Naine”), which he called G. falcatus var. ternatensis (De Man).

The identity of G. bossorotundus Roxas and Estampador, 1930, found in a collection of G. falcatus sensu lato, from the Philippines, cannot be determined from their account; it appears to be a member of the Falcatus group with a damaged telson. According to E. Estampador (pers. comm.), the unique holotype was lost during World War II, so the problem may never be resolved.

While studying the biology of stomatopods in Thailand in 1974, H. Dingle, University of Iowa, and R. Caldwell, University of California, recognized that two distinct species of Gonodactylus, both identifiable on morphological grounds with G. falcatus, occurred there. Representatives of the two forms were sent to me for identification, and, in order to provide those identifications, I had to reexamine available types of species in the Falcatus group. As a result, I recognized five species of that group in addition to G. graphurus: G. falcatus (Forskål, 1775), G. glabrous Brooks, 1886, G. mutatus Lanchester, 1903, G. ternatensis De Man, 1902, and a new species, G. randalli.

In their study of two species of the Falcatus group from Thailand, Dingle and Caldwell noted differences in color between males and females of the same species, and differences in color, size, and biology between two species living on the same tidal flat. This kind of information is badly needed for all members of the Falcatus group. Study series documented with information on habitat and color in life would be most welcome.

Species recognition in Gonodactylus is hampered by the paucity of clearly defined characters in members of this genus, coupled with the masking effects of aging as well as secondary sexual changes, especially in males. Although most stomatopods can be identified through meristic features, in Gonodactylus species discrimination is based largely on subjective features, including: relative length of the apical spine and shape of the rostral plate, size and shape of the ocular scales, ornamentation and shape of the telson, and structure and armature of the uropods. Rarely are there such clear, distinctive characters as the transverse grooves on the abdomen of G. graphurus or the accessory carina of the telson in the American G. oerstedii Hansen.

Representatives of the Falcatus group have been cited in the literature more than 100 times, and since few records include descriptive notes or figures, virtually none of them can be identified with certainty without reexamination of the material cited. For this reason and because synonymies are being compiled by L. B. Holthuis, Rijksmuseum van Natuurlijke Historie, Leiden, for the Crustaceorum Catalogus, I have included here only the original citation for each species and selected subsequent citations. Much more material will have to be studied before the ranges, synonymies, and extent of morphological and geographical variation of these species can be worked out. The accounts given here have been restricted to descriptions, illustrations of types of four previously named species, illustrations of topotypic material for one species, and the account of a new species.

Additional species in the Falcatus group may be recognized in the future. A. Michel (in litt.), Centre National pour l'Exploitation des Oceans, Tahiti, has been studying stomatopod larvae from the central Pacific for several years and can recognize the larvae of six separate species of the Falcatus group of Gonodactylus occurring in that area.

Further, with our current state of knowledge, not all of the material of the Falcatus group available for study can be identified with certainty. The population of this group from Hawaii, identified with G. falcatus and thought to be a recent introduction there (Kinzie, 1968), cannot now be identified with any of the species described below. Kinzie (1968, fig. 3) has shown the sixth abdominal somite and telson of a male and a female from Hawaii: the female has a sharp median carinule on the sixth abdominal somite, but the male lacks it. This carinule is present in both sexes of G. faicatus from the Red Sea and, indeed, is characteristic of only G. falcatus, G. graphurus, and G. randalli, new species, in the Falcatus group. This carinule is not apparent in the limited material from Hawaii in the collection of the National Museum of Natural History. This may prove to be a variable feature of G. falcatus, but such variation is not found in topotypic material of that species.

Serène (1954:82) pointed out that five morphological changes accompanied aging in G. mutatus (as G. falcatus var. ternatensis) from Viet Nam, and his comments probably are applicable to members of the Falcatus group in general: (a) thickening of the margins of the telson, resulting in blunter teeth with less space between them; (b) comparative inflation of the carinae of the sixth abdominal somite and the telson, relatively accentuated in males; (c) enlargement of the intermediate tooth of the telson; (d) reduction of the number of submedian marginal denticles on the telson; and (e) reduction in size and number of posterior spines on the carinae of the sixth abdominal somite and telson. Serène also illustrated differences in rostral plates of males and females (fig. 13) and telsons (pls. 9, 10). All of these changes must be taken into account in any study of members of the Falcatus group.

Because such changes are less drastic in females than in males, figures given herein are, with one exception, based on adult females of similar size. Females show less change with age than do males, especially in inflation of the carinae of the sixth abdominal somite and telson. In adult males the inflation of these carinae may completely mask other morphological features. Figure 5, based on a male of G. rotundus Borradaile, shows the extent of inflation of the carinae of the telson in males of that species. The median carina of the telson is so inflated in that specimen that the knob, clearly bilobed in juveniles and females, is scarcely or not at all discernible.

In addition to the traditional illustrations included in the species accounts below, comparative outline drawings in a stylized format also are presented for the five species reported here (Figures 9–Figures 13). The format is designed after that used very effectively by Hobbs (1974) for North American crayfishes, and will be used in the future to supplement species accounts whenever possible. Eventually I hope to prepare an identification manual for the Indo-West-Pacific stomatopods using this technique.

I have not included a key to the species of the Falcatus group, even though such a key could easily be constructed for the six species of the complex now recognized. In view of the uncertain status of some of the population of the Falcatus group, I believe that a key would be more misleading than helpful at this time.

Terms and measurements used in the accounts below have been explained in earlier papers (Manning, 1966, 1968). All measurements are in millimeters (mm).

The institutions in which the material reported here is deposited are identified by the following acronyms:

BMNH

British Museum (Natural History), London

MZC

University Museum of Zoology, Cambridge, England

SMF

Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt am Main, West Germany

USNM

National Museum of Natural History, Smithsonian Institution, Washington (formerly United States National Museum)

ACKNOWLEDGMENTS.—H. Dingle, Department of Zoology, University of Iowa, and R. Caldwell, Department of Zoology, University of California, Berkeley, initiated this study by forwarding material for identification and further helped by sharing their field observations on these and other species of somatopods. Marjorie Reaka, Department of Zoology, University of Maryland, also discussed various aspects of the biology of stomatopods based on her field observations. A. Michel, Centre National pour l'Exploitation des Oceans, Tahiti, commented on his studies on stomatopod larvae, forwarded copies of illustrations of larvae, and loaned specimens. At the Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt am Main, West Germany, the late R. Bott provided working space and accommodations in 1971 and, more recently, M. Türkay loaned the syntypes of G. ternatensis De Man. R. W. Ingle, Department of Zoology, British Museum (Natural History), London, loaned the holotype of G. glabrous Brooks. C. B. Goodhart, University Museum of Zoology, Cambridge, England, loaned the syntypes of G. mutatus Lanchester and G. rotundus Borradaile. The illustrations were prepared by my wife Lilly. Anne Cohen and Cynthia Hemming aided in proofreading and checking references and L. S. Kornicker reviewed the manuscript.

Part of this study was supported by the Smithsonian Institution through its Research Awards Program.

Gonodactylus falcatus (Forskål, 1775)

FIGURES 1, 2a, 9

Cancer falcatus Forskål, 1775:96.

Gonodactylus falcatus.—Holthuis, 1967:31, 41 [complete synonymy for Red Sea].

MATERIAL.—SW tip of Umm Aabak, N of Nocra, Dahlak Archipelago, Eritrea, Ethiopia, Red Sea; 0-2 m deep, among corals, sandy bottom; Israel South Red Sea Expedition no. 1447; 5–6 Apr 1962: 4art, TL (total length) 47–62 mm; 7art, TL 46–59 mm (USNM 119278).

DESCRIPTION.—Rostral plate (Figures 1a, 9c) as long as broad or slightly broader than long, median spine relatively short; anterior margins of plate straight, perpendicular to body line or sloping posteriorly; anterolateral angles of plate acute but broadly rounded. Ocular scales (Figures 1a, 9c) small, breadth of both about one-fourth plate width, erect, set transversely to body line, apices rounded. Anterior 5 abdominal somites (Figure 2a) lacking transverse grooves, transverse rows of light black spots present in some specimens. Sixth abdominal somite (Figures 1b, 9a) with sharp median carinule and with 6 dorsal carinae, variously inflated, usually armed posteriorly; apical spinules often obsolete, especially in adult males; submedian and intermediate carinae flattened dorsally, with posterior excavation under each spine. Pair of small black spots (usually visible in preserved specimens) present anteriorly between submedian and intermediate carinae. Telson (Figures 1b, 9a) with length and width subequal or width slightly greater, appearing broad. Dorsal carinae of telson usually sharp, more inflated in adult males, median and accessory medians each usually with apical spinule flanked ventrally by rounded excavation. Knob strongly bilobed. Three pairs of marginal teeth present, submedians broad, with movable apices, intermediates broader, apices sharp, laterals short, apices sharp. Well-marked black spot present on anterior surface anterior to each dorsal submedian carina. Ventral surface of telson with low postanal ridge and longer ventral carina on each submedian tooth. Uropodal endopod with single line of marginal setae, those on outer margin directed dorsally. Basal prolongation of uropod (Figures 1d, 9d) with 1 lobe proximally on inner margin of outer spine.

FIGURE 1.—Gonodactylus falcatus (Forskål), female, TL 50 mm, Red Sea: a, rostral plate and ocular scales; b, sixth abdominal somite and telson, dorsal view; c, telson, lateral view; d, basal prolongation of uropod, ventral view.

MEASUREMENTS.—Males (4), TL 47–62 mm; females (7), TL 46–59 mm. Holthuis (1967) recorded males as large as 70 mm and females as large as 66 mm from the Red Sea. Other measurements of female, TL 59 mm: carapace length 13.0; rostral plate length 3.6, width 3.8; fifth abdominal somite width 10.0; telson length 8.3, width 8.5.

REMARKS.—This account of Gonodactylus falcatus is based on material from the Red Sea, the type-locality. Forskål's types were not traced; they probably are not extant. Although this appears to be the common species of the Falcatus group in the Red Sea, it is known now that G. mutatus also occurs there, as discussed below in the account of that species.

Gonodactylus falcatus is a distinctive species, one of three in the Falcatus group with a sharp median carinule on the sixth abdominal somite; the carinule is present in all of the material I have examined of this species from the Red Sea. The median carinule on the sixth abdominal somite is present also in G. graphurus and G. randalli, new species, described below. Gonodactylus falcatus differs from G. graphurus in numerous features, the most important of which is the absence of fine, transverse grooves on the abdominal somites, as noted by several authors, including Kemp (1913). Sketches of the abdomen of both species are shown in Figure 2. Differences between G. falcatus and G. randalli are discussed under the account of the latter species.

Both G. glabrous and G. mutatus lack the median carinule on the sixth abdominal somite, but both resemble G. falcatus and differ from G. ternatensis in having a distinctly bilobed knob as well as in having a relatively short apical spine on the rostral plate (Figures 9–13). The dorsal carinae of the telson of G. falcatus are not so sharp as those of G. glabrous and are never so tumid as in adult males of G. mutatus (Figure 5).

Gonodactylus falcatus, like G. ternatensis, is a relatively large species of Gonodactylus, attaining a comparatively much greater size than does G. mutatus (TL 60 mm or less adults) and perhaps G. glabrous and G. randalli as well.

FIGURE 2.—Abdomen, lateral view: a, Gonodactylus falcatus (Forskål); b, Gonodactylus graphurus Miers.

DISTRIBUTION.—Known with certainty from the Red Sea; Holthuis (1967) has summarized all records from that area. Its distribution outside of the Red Sea remains to be determined. Despite persistent reports of the occurrence of G. falcatus in the Mediterranean, there are no verified records of adults from that sea (Manning, in press).

Gonodactylus glabrous Brooks, 1886

FIGURES 3, 10

Gonodactylus glabrous Brooks, 1886:22, 62, pl. 14: fig. 5, pl. 15: figs. 7, 9.

Gonodactylus glabrous var. ternatensis De Man, 1902:914, pl. 27: fig. 67 [part, figure of female only].

MATERIAL.—Samboangan Reefs, Philippine Islands; Challenger, leg.; holotype: 1 art, TL 50 mm (BMNH).—Ternate, Molucca Islands, Indonesia; Kükenthal, leg.; 1894; paralectotype of G. glabrous var. ternatensis De Man: 1 art, TL 51 mm (ex SMF 5779).

DESCRIPTION.—Rostral plate (Figures 3a, 10c) as long as broad or slightly broader than long, median spine appearing short; anterior margins of plate sloping anteriorly; anterolateral angles of plate acute but rounded, appearing angled. Ocular scales (Figures 3a, 10c) small, breadth of both slightly less than one-third rostral plate width, erect, set transversely to body line, apices rounded dorsally. Anterior 5 abdominal somites lacking transverse grooves but each with transverse rows of dark spots (at least in females). Sixth abdominal somite (Figures 3b, 10a) lacking median carinule, with 6 short, sharp carinate spines, carinae of submedian and intermediate spines flattened dorsally. Pair of small black spots not present anteriorly on sixth somite (not visible on either specimen examined). Telson (Figures 3b, 10a) broader than long, appearing slender. Dorsal carinae of telson sharp, median with long apical spine and accessory medians each with short apical spine, apex of each flanked ventrally by rounded excavation. Knob bilobed, emarginate in midline. Three pairs of marginal teeth present, submedians slender, with movable apices, intermediates broader, apices sharp, laterals short, sharp. Anterior part of telson lacking black spot anterior to each dorsal submedian carina. Ventral surface of telson with postanal ridge and long, sharp ventral carina on each submedian tooth. Uropodal endopod with single line of marginal setae, those on outer margin directed dorsally. Basal prolongation of uropod (Figures 3d, 10d) with 1 lobe proximally on inner margin of outer spine.

FIGURE 3.—Gonodactylus glabrous Brooks, female holotype, TL 50 mm, Philippines: a, rostral plate and ocular scales; b, sixth abdominal somite and telson; c, telson, lateral view; d, basal prolongation of uropod, ventral view.

MEASUREMENTS.—Only females (2) examined, TL 50–51 mm. Other measurements of larger specimen: carapace length 11.8; rostral plate length 3.6, width 3.8; fifth abdominal somite width 10.0; telson length 7.7, width 8.2.

REMARKS.—I have examined only two specimens of this species, which I believe should be recognized as distinct from the others in the Falcatus group. Gonodactylus glabrous lacks the median carinule on the sixth abdominal somite that is found in G. falcatus and G. randalli, new species (described below), and is larger and has much sharper carinae on the telson in adults than do specimens of G. mutatus. Gonodactylus glabrous differs from G. ternatensis in having a bilobed rather than single knob, and a much shorter spine on the rostral plate. Gonodactylus glabrous resembles these species and differs from G. graphurus in lacking transverse grooves on the abdomen.

Paired black spots on the sixth abdominal somite and on the anterior part of the telson are characteristic of most species of the Falcatus group. Even if they are not visible in preserved material, their position often is indicated by distinct, shallow pits. Gonodactylus glabrous may be ornamented with transverse lines of dark spots on the anterior five abdominal somites rather than the paired spots on the sixth abdominal somite and telson. Brooks (1886) clearly showed and described the transverse rows on the anterior abdominal somites, and De Man (1902) commented on the occurrence of similar rows of spots on the abdomen of the female of his G. ternatensis, identified here with G. glabrous. He also noted the absence of these anterior abdominal spots but remarked on the presence of the paired spots on the sixth abdominal somite and telson in his other specimen, that selected here as the lectotype of G. ternatensis. Inasmuch as De Man's specimens came from the same collection, it seems unlikely that the observed differences in the position of the spots are the results of preservation techniques.

DISTRIBUTION.—Known with certainty from the Philippines and from the Molucca Islands; its depth range and habitat preferences are unknown.

Gonodactylus mutatus Lanchester, 1903

FIGURES 4, 5, 11

Gonodactylus chiragra var. mutatus Lanchester, 1903:450.

Gonodactylus glaber var. rotundus Borradaile, 1907:211 [key], 212, pl. 22: fig. 2.—Dollfus, 1938, fig. 20.

Gonodactylus falcatus var. ternatensis.—Serène, 1954:6, 7, 10, 74, 48, 80, fig. 13–7, 8, pl. 4: figs. 7–12, pl. 10 [not Gonodactylus ternatensis De Man, 1902].

MATERIAL.—Phuket, Thailand; H. Dingle and R. L. Caldwell, leg.; Jun-Aug 1974: 7 art, TL 21–33 mm; 14 art, TL 24–41 mm (USNM 168516). Furnadu Velu, Miladumadulu Atoll, Maldive Islands; J. S. Gardiner, leg.; lectotype of G. mutatus: 1 art, TL 44 mm (MZC). Data same; paralectotype of G. mutatus: 1 art, TL 38 mm (MZC). Goidu, Goifurfehendu Atoll, Maldive Islands; J. S. Gardiner, leg.; paralectotypes of G. mutatus: 2 art, TL 24–32 mm; 1 art, TL 34 mm (MZC). Hulule, Male Atoll, Maldive Islands; J. S. Gardiner, leg.; paralectotype of G. mutatus: 1 art, TL 22 mm (MZC). Coetivy, Seychelles Islands; J. S. Gardiner, M. A. Caius, leg.; syntype of G. rotundus: 1 art, TL 44.5 mm (MZC). Zanzibar; C. Crossland, leg.; 1901; syntype of G. rotundus: 1 art, TL 36 mm (MZC).

An additional lot of syntypes of G. rotundus from Coetivy was examined; the specimen(s) have completely disintegrated.

DESCRIPTION.—Rostral plate (Figures 4a, 5a, 11c) with length and width subequal or width greater, median spine relatively short; anterior margins of plate straight, direction of slope variable; anterolateral angles acute but broadly rounded. Ocular scales (Figures 4a, 11c) small, breadth of both about one-third rostral plate width, erect, set transversely to body line, apices rounded dorsally. Anterior 5 somites lacking transverse grooves or rows of dark spots. Sixth abdominal somite (Figures 4b, 5b, 11c) lacking median carinule, with 6 variously inflated carinae, each usually with posterior spine; spines of submedian and intermediate carinae often missing in large specimens of both sexes. Pair of small black spots present anteriorly on sixth somite, each set lateral to submedian carina. Telson Figures 4b, 5b, 11c) with length and width subequal or width greater, appearing broad. Dorsal carinae of telson variously inflated, tumid in adults of both sexes, especially so in males, median and accessory medians each with posterior spinule, often missing in large specimens of both sexes; dorsal carinae may be so inflated that they almost merge (Figure 5). Knob bilobed medially emarginate (indistinct in large males). Three pairs of broad marginal teeth present, submedians with movable apices, laterals bluntly pointed. Well-marked black spot present anterior to each dorsal submedian carina. Ventral surface of telson with obscure postanal ridge and longer carina on each submedian tooth. Uropodal endopod with single line of marginal setae, those on outer margin directed dorsally. Basal prolongation of uropod (Figures 4d, 11d) with 1 lobe proximally on inner margin of outer spine.

MEASUREMENTS.—Males (13), TL 21–44.5 mm; females (16), TL 24–44 mm. The largest specimen recorded by Seréne (1954) was a female 57 mm long; he noted that representatives of this species rarely attained 60 mm in length. Other measurements of female lectotype, TL 44 mm: carapace length 9.2; rostral plate length 2.8, width 2.8; fifth abdominal somite width 7.7; telson length 5.4, width 6.5.

REMARKS.—Gonodactylus mutatus is a relatively small species of Gonodactylus in which the dorsal carinae of the telson tend to be very tumid. The tumidity of these carinae was noticed and figured by Seréne (1954, pl. 10: fig. 5) and was one of the characteristics stressed by Borradaile (1907) in his account of G. rotundus. Lanchester (1903) did not illustrate his material of G. mutatus, but the female lectotype is illustrated here (Figure 4). The telson of a male syntype of G. rotundus, here considered to be a synonym of G. mutatus, also is shown here (Figure 5) to show the difference in tumidity of the telson carinae in the two sexes. The knob is indistinctly bilobed in adult males as shown in Figure 5. The telson of a male, TL 47 mm, from the Red Sea was illustrated by Dollfus (1938, fig. 20); it resembles that shown here in Figure 5.

FIGURE 4.—Gonodactylus mutatus Lanchester, female lectotype, TL 44 mm, Maldive Islands: a, rostral plate and ocular scales; b, sixth abdominal somite and telson; c, telson, lateral view; d, basal prolongation of uropod, ventral view.

I can find no major differences between representatives of this species from the western Indian Ocean and the single lot available from Thailand. The anterior margins of the rostral plate tend to slope forward in the eastern specimens (see Serène, 1954, fig. 13–7,8) and are perpendicular to the body line or slope posteriorly in the western specimens. The dorsal carinae of the telson appear to be slightly less tumid in eastern specimens. Geographic variation in such features is not known at this time for any species of Gonodactylus.

This species agrees with G. glabrous and G. ternatensis and differs from G. falcatus, G. graphurus, and G. randalli in lacking the median carinule on the sixth abdominal somite. It differs from G. ternatensis in having a much shorter spine on the rostral plate, a much slenderer telson, and a bilobed knob; in addition, it is a much smaller species. Gonodactylus mutatus resembles G. glabrous in many respects, but in the latter species the carinae of the telson are quite sharp even in adult females (the males are unknown) and the rostral plate is more angled anterolaterally.

FIGURE 5.—Gonodactylus mutatus Lanchester, male, TL 44.5 mm. syntype of G. rotundus Borradaile, Coetivy, Seychelles Islands: a, rostral plate; b, sixth abdominal somite and telson.

The largest specimen from Lanchester's type-series, a female 44 mm long from Furnadu Velu, is here selected as the lectotype of G. mutatus.

DISTRIBUTION.—Indo-West-Pacific region. It is known with certainty from Viet Nam, Thailand, the Maldive Islands, the Seychelles Islands, Zanzibar, and from one locality in the Red Sea. The extent of its distribution within the region remains to be determined. It apparently lives in shallow water, on coral or coral rubble.

Gonodactylus randalli, new species

FIGURES 6, 12

MATERIAL.—Arno Atoll, Marshall Islands; R. W. Hiatt, leg.; sta E2–377; 1950; paratype: 1 art, TL 20 mm (USNM 124365). Canton Island, Phoenix Islands; from coral head in lagoon; 20–25 ft (6–8 m); C. A. Ely, leg.; 2 Nov 1941; paratype: 1 art, TL 39 mm (USNM 124362). Outside barrier reef, 200 yds (ca. 180 m) E of pass of Papetoai Bay, Moorea, Society Islands; 70 ft; J. E. Randall, leg.; 15 May 1957; holotype: 1 art, TL 51 mm (USNM 124372).

DESCRIPTION.—Rostral plate (Figures 6a, 12c) with length and width subequal or length slightly greater, median spine appearing short; anterior margins of plate almost straight, perpendicular to body line; anterolateral angles of plate broadly rounded. Ocular scales (Figures 6a, 12c) very small, erect, apices rounded, width of scales about one-third plate width. Anterior abdominal somites lacking transverse grooves and transverse rows of dark spots. Sixth abdominal somite (Figures 6b, 12a) with prominent median carinule and 6 carinate spines, submedians and intermediates distinctly flattened dorsally. Pair of small black spots present anteriorly on sixth somite (on 1 paratype, not visible in others), each spot situated lateral to submedian carina. Telson (Figures 6b, 12a) appearing elongate, length and width subequal or width slightly greater. Dorsal carinae of telson sharp, slender, median and accessory medians each with long apical spine, each spine flanked ventrally by carinate excavation. Knob indistinctly bilobed in holotype, distinctly bilobed in paratypes. Three pairs of sharp, slender marginal teeth present, submedians with movable apices, apices of intermediates and laterals spiniform (left lateral tooth missing in holotype). Anterior margin of telson with pair of black spots (on 1 paratype), one lateral to each dorsal submedian carina. Ventral surface of telson with sharp postanal ridge and sharp longitudinal carina on each submedian tooth. Uropodal endopod with multiple rows of setae, some of those on lateral margin directed dorsally. Basal prolongation of uropod (Figures 6d, 12d) with 2 proximal lobes between spines.

MEASUREMENTS.—Only females (3) known, TL 29–51 mm. Other measurements of female holotype, TL 51 mm; carapace length 10.9; rostral plate length 3.4, width 3.4; fifth abdominal somite width 8.6; telson length 6.8, width 7.0.

FIGURE 6.—Gonodactylus randalli, new species, female holotype, TL 51 mm, Moorea: a, rostral plate and ocular scales; b, sixth abdominal somite and telson (left lateral tooth missing); c, telson, lateral view; d, basal prolongation of uropod, ventral view.

REMARKS.—Gonodactylus randalli, like G. falcatus and G. graphurus, has a strong median carinule on the sixth abdominal somite. That carinule, the short, broad rostral plate, the presence of 2 distinct lobes between the spines of the basal prolongation of the uropod, and the multiple marginal setae on the uropodal endopod distinguish G. randalli from all of the species of the Falcatus group that lack the transverse grooves on the abdomen.

One of the paratypes shows the dark spots on the telson that are characteristic of most species of the Falcatus group; these are not visible in the other specimens although their position is marked by faint, unpigmented pits. The spots are not shown in Figure 6b, based on the holotype.

ETYMOLOGY.—This species is named for the collector of the holotype, John E. Randall.

DISTRIBUTION.—Pacific Ocean, from the Marshall Islands, the Phoenix Islands, and the Society Islands, in depths between 20 and 70 ft (12–21 m).

Gonodactylus ternatensis De Man, 1902

FIGURES 7, 8, 13

Gonodactylus glabrous var. ternatensis De Man, 1902:914 [part; not figured specimen].

Gonodactylus glabrous.—Bigelow, 1931:127, fig. 1 [part; specimens from Samoa?] [not Gonodactylus glabrous Brooks, 1886].

Gonodactylus falcatus.—Serène, 1954:6, 79, fig. 13–6, pl. 9 [not Gonodactylus falcatus (Forskål, 1775)].

MATERIAL.—Apia, Upolu Island, Samoa Islands; D. S. Jordan, leg.; Jul 1902: 1art, TL 50 mm (USNM 64853). Phuket, Thailand; H. Dingle and R. Caldwell, leg.; 1974: 5art, TL 43–68 mm; 9 art, TL 23–66 mm (USNM 156252). Ternate, Molucca Islands, Indonesia; Kükenthal, leg.; 1894; lectotype: 1 art, TL 56.5 mm (SMF 5779).

DESCRIPTION.—Rostral plate (Figures 8a, 13a) longer than broad, median spine relatively long; anterior margins of plate sloping anteriorly; anterolateral angles of plate acute but rounded. Ocular scales (Figures 8a, 13a) small, breadth of both about one-third rostral plate width, erect, set obliquely to body line, apices rounded or slightly flattened dorsally. Anterior abdominal somites lacking transverse grooves and rows of dark spots. Sixth abdominal somite (Figures 7, 8b, 13a) lacking median carinule, with 6 long, sharp, carinate spines, carinae of submedian and intermediate spines flattened dorsally. Pair of small black spots present anteriorly on sixth somite, each situated lateral to submedian carina (faded in illustrated specimen). Telson (Figures 7, 8b, 13a) longer than broad, appearing elongate. Dorsal carinae of telson sharp, median and accessory medians each with long apical spine flanked ventrally by rounded excavation. Knob single, undivided. Three pairs of long, sharp marginal teeth present, submedians with movable apices, laterals well developed. Black spot present on anterior surface of telson lateral to each dorsal submedian carina (faded on illustrated specimen). Ventral surface of telson with postanal keel (reduced in some specimens) and long, sharp ventral carina on each submedian tooth. Uropodal endopod with single line of marginal setae, those on outer margin directed dorsally. Basal prolongation of uropod (Figures 8d, 13d) with 1 lobe proximally on inner margin of outer spine.

FIGURE 7.—Gonodactylus ternatensis De Man, female lectotype, TL 56.5 mm, Ternate: sixth abdominal somite and telson.

MEASUREMENTS.—Males (5), TL 43-68 mm; females (11), TL 23-66. Other measurements of female lectotype, TL 56.5 mm: carapace length 13.5; rostral plate length 4.4, width 4.2; fifth abdominal somite width 11.4; telson length 10.2, width 9.3.

REMARKS.—Gonodactylus ternatensis is a relatively large species of Gonodactylus, and, like G. graphurus, can be recognized readily in preserved material. The elongate rostral plate, with an anterior spine comparatively much longer than in any other species of the Falcatus group, the long telson, with sharply spined longitudinal carinae, and the single, undivided knob are all characteristic of the species.

The larger of the two specimens referred to this species by De Man, a female TL 56.5 mm, is here selected as the lectotype. The smaller female, the paralectotype, is identified with G. glabrous herein. There is little doubt that G. ternatensis would have been recognized sooner if De Man had illustrated his specimen of G. ternatensis sensu stricto rather than the G. glabrous in his original account.

It seems likely that most or all of the specimens reported by Bigelow (1931) from Samoa as G. glabrous actually are G. ternatensis. When Bigelow compared his material from Samoa with material from the Philippines (probably based on materials collected there by the Albatross, which then were being studied by Bigelow), he noted (p. 130): “In the Samoan specimens the maximum length of the rostrum is generally considerably greater than the width at anterolateral angles. The anterior margins on each side of the spine are nearly transverse and the anterolateral angles are broadly rounded. In the Philippine specimens the length and breadth are about equal, the anterior margins are usually directed obliquely forward from the junction with the spine and the anterolateral angles are narrowly rounded or subacute.” The anterolateral angles of the rostral plate are more acute in the two specimens of G. glabrous that I have examined than they are in G. ternatensis.

The species from Viet Nam identified with G. falcatus by Seréne (1954) is, as shown by his figures of the rostral plates and telsons, G. ternatensis sensu stricto, whereas his material identified with G. falcatus var. ternatensis has been referred to G. mutatus herein.

FIGURE 8.—Gonodactylus ternatensis De Man, female, TL 50 mm, Samoa: a, rostral plate and ocular scales; b, sixth abdominal somite and telson; c, telson, lateral view; d, basal prolongation of uropod, ventral view.

DISTRIBUTION.—This species is known with certainty from Apia, Samoa; Phuket, Thailand; Viet Nam; and Ternate, Molucca Islands, Indonesia. Apparently G. ternatensis is a shallow-water species, living near shore on flats exposed at low tide.

FIGURES 9–13.—Five species of the Falcatus group of Gonodactylus: 9, G. falcatus (Forskål); 10, G. glabrous Brooks; 11, G. mutatus Lanchester; 12, G. randalli, new species; 13, G. ternatensis De Man. (Figures 9–13: a, sixth abdominal somite and telson; b, apices of submedian teeth of telson, ventral view; c, rostral plate and ocular scales; d, basal prolongation of uropod.)

Literature Cited

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Borradaile, L. A.

1907.   Stomatopoda from the Western Indian Ocean. In The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the Leadership of Mr. J. Stanley Gardiner. Transactions of the Linnaean Society of London, (2)12:209–216, plate 22.

Brooks, W. K.

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Dollfus, R. Ph.

1938.   Stomatopoda, II: Catalogue Synonymique des Espèces Jusqu'à Présent Récoltées dans la Mer Rouge, y Compris la Partie Sud du Canal de Suez et le Golfe d'Aden. In Mission Robert Ph. Dollfus en Égypte, X. Mémoires de l'Institut d'Égypte, 37:185–236, figures 1–8, 10, 11, 13–23.

Fabricius, J. C.

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Hansen, H. J.

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1971.   On the Nomenclature of Gonodactylus graphurus White, 1847 (Nomen Nudum), Miers, 1875 (Stomatopoda, Gonodactylidae). Crustaceana, 21(2):220–221.

Kemp, S.

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Kinzie, Robert A., III

1968.   The Ecology of the Replacement of Pseudosquilla ciliata (Fabricius) by Gonodactylus falcatus (Forskål) (Crustacea; Stomatopoda) Recently Introduced into the Hawaiian Islands. Pacific Science, 22(4):465–475, figures 1–5.

Lanchester, W. F.

1903.   Stomatopoda, with an Account of the Varieties of Gonodactylus chiragra. Marine Crustaceans, VIII. Pages 444–459 in part 4 of volume 1 in J. S. Gardiner, The Fauna and Geography of the Maldive and Laccadive Archipelagoes, Being the Account of the Work Carried on and of the Collections Made by an Expedition during the Years 1899 and 1900. Plate 23.

De Man, J. G.

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Manning, Raymond B.

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