New and Rare Stomatopod Crustacea from the Indo-West-Pacific Region RAYMOND B. MANNING ' SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 264 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoo/ogy Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world cf science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. S. Dillon Ripley Secretary Smithsonian Institution S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 2 6 4 New and Rare Stomatopod Crustacea from the Indo-West-Pacific Region Raymond B. Manning SMITHSONIAN INSTITUTION PRESS City of Washington 1978 A B S T R A C T Manning, Raymond B. New and Rare Stomatopod Crustacea from the Indo- West-Pacific Region. Smithsonian Contributions to Zoology, number 264, 36 pages, 16 figures, 1978.?Nineteen species of stomatopod crustaceans are reported. Five new genera are recognized: Hoplosquilloides, for H. coronatus, new species: Keppelius, for Lysiosquilla hystricotelson Barnard, 1958; Pullosquilla, for Aus- trosquilla litoralis Michel and Manning, 1971, A. malayensis Manning, 1968, and a new species, P. thomassini; Busquilla, for Squilla quadraticauda Fukuda, 1911, and B. plantei, new species; and Tuleariosquilla, for T. parvula, new species. Five other new species are described: Alima orientalis, Clorida japonica, Cloridopsis aquilonaris, Harpiosquilla malagasiensis, and Neocoronida martensi. Four species are redescribed: Clorida minor (Jurich, 1904); Gonodactylus viridis Serene, 1954; Neocoronida trachurus (von Martens, 1881); and Parvisquilla multituberculata (Borradaile, 1898). Gonodactylus confinis de Man, 1902, and G. segregatus Lanchester, 1903 are shown to be synonyms of G. affinis de Man, 1902; Coronis spinosa Wood-Mason, 1875 is a synonym of Coronis tricarinata Claus, 1871; and Parvisquilla xishaensis Liu, 1975 is a synonym of Parvisquilla multituberculata (Borradaile, 1898). Additional material of Clorida latispina Manning, 1968 and C. malaccensis Manning, 1968 is reported; C. malaccensis var. moluccensis Moosa, 1973 is a synonym of the latter. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is re- corded in the Instftmion's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavemosa (Linnaeus). Library of Congress Cauloging in Publication Data Manning, Raymond B 1934- New and rare stomatopod Crustacea from the Indo-West-Pacific region. (Smithsonian contributions to zoology ; 264) Bibliography: p. 1. Stomatopoda. 2. Rare animals?Indian Ocean. 3. Rare animals?Pacific Ocean. 4. Crus- tacea?Classification. 5. Crustacea?Indian Ocean. 6. Crustacea?Pacific Ocean. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; 264. QL1.S54 no. 264 [QL444.M.375] 591'.08s [595'.382'09165] 77-14228 Contents Page Introduction 1 Acknowledgments 1 Family GONODACTYLIDAE Giesbrecht, 1910 2 Genus Gonodactylus Berthold, 1827 2 Gonodactylus affinis de Man, 1902 2 Gonodactylus viridis Serene, 1954 4 Hoplosquilloides, new genus 6 Hoplosquilloides coronatus, new species 6 Family LYSIOSQUILLIDAE Giesbrecht, 1910 8 Genus Heterosquilla Manning, 1963 8 Heterosquilla tricarinata (Claus, 1871) 8 Keppelius, new genus 9 Keppelius hystricotelson (Barnard, 1958) 10 Genus Neocoronida Manning, 1976 11 Key to Species of Neocoronida 11 Neocoronida martensi, new species 12 Neocoronida trachurus (von Martens, 1881) 13 Genus Parvisquilla Manning, 1973 15 Key to Species of Parvisquilla 16 Parvisquilla multituberculata (Borradaile, 1898) 16 Pullosquilla, new genus 18 Key to Species of Pullosquilla 19 Pullosquilla litoralis (Michel and Manning, 1971) 19 Pullosquilla thomassini, new species 20 Family SQUILLIDAE Latreille, 1803 21 Genus Alima Leach, 1817 21 Alima orientalis, new species 21 Busquilla, new genus 23 Busquilla plantei, new species 23 Genus Clorida Eydoux and Souleyet, 1842 25 Clorida japonica, new species 25 Clorida latispina Manning, 1968 26 Clorida malaccensis Manning, 1968 26 Clorida minor (Jurich, 1904) 27 Genus Cloridopsis Manning, 1968 28 Cloridopsis aquilonaris, new species 28 Genus Harpiosquilla Holthius, 1964 30 Harpiosquilla malagasiensis, new species 30 Tuleariosquilla, new genus 30 Tuleariosquilla parvula, new species 32 Literature Cited 34 New and Rare Stomatopod Crustacea from the Indo-West-Pacific Region Raymond B. Manning Introduction Visits to European museums in 1971 and to the Zoological Survey of India, Calcutta in 1972, and examination of collections received since then have resulted in the accumulation of a large amount of new information on Indo-West-Pacific stomatopods. Because some names are required by colleagues working in other fields, redescription of five known species, observations on types of several poorly known species, and descriptions of new taxa, in- cluding five new genera and nine new species, are presented here. More cohesive summaries of other Indo-West-Pacific stomatopods, at the generic or species-group levels are in preparation or in press; these include reviews of Lysiosquilla and the mem- bers of the falcatus group of Gonodactylus and redescriptions of types and descriptions of new spe- cies in Oratosquilla. A general review of stomato- pod taxa at and above the generic level is also in preparation. All measurements are in millimeters (mm). TL refers to total length, measured on the midline from the anterior margin of the rostral plate to the apices of the submedian teeth of the telson; CL refers to carapace length, measured on the mid- line, exclusive of rostral plate. Terms and measure- Raymond B. Manning, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C. 20560. ments have been explained in earlier papers (see Manning, 1969a). The descriptive accounts vary in complexity de- pending on the taxon involved; thus a diagnosis is given for a new Alima and a new Cloridopsis, genera in which relatively few distinguishing char- acteristics are known. Often, characters mentioned in generic diagnoses are not repeated in descrip- tions of species in that genus. The following abbreviations are used to designate repositories: BMN'H British Museum (Natural History), London MXHXP Museum National d'Histoire Naturelle, Paris MRAC Musee Royal de 1'Afrique Centrale, Tervuren, Belgium MZC University Museum of Zoology, Cambridge, Eng- land SME Station Marine d'Endoume, Marseille SMF Natur-Museum und Forschungsinstitut Seiicken- berg, Frankfurt am Main USNM National Museum of Natural History, Smithso- nian Institution, Washington (USNM = former United States National Museum) ZMB Zoologisches Museum an der Humboldt-Universi- ta't zu Berlin ZMH Zoologisches Institut und Zoologisches Museum, Hamburg ZSI Zoological Survey of India, Calcutta ZSM Zoologisches Staatsammlung, Munich ACKNOWLEDGMENTS.?A large part of this report was made possible by B. Thomassin, M. Peyrot- Clausade, and R. Plante, Station Marine d'En- doume, Marseille, all of whom sent samples of unidentified stomatopods, largely from Madagascar, SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY for identification. H. Dingle, Department of Zool- ogy, University of Iowa, and R. Caldwell, Depart- ment of Zoology, University of California, Berke- ley, provided material from their field studies of stomatopods in Thailand and freely discussed their field observations. The following individuals also provided working space and/or material for study on loan at various times: P.L.G. Benoit, Musee Royal de l'Afrique Centrale, Tervuren (directly and indirectly through L. B. Holthuis, Rijks- museum van Natuurlijke Historic Leiden); H. Fechter, Zoologisches Staatsammlung, Munich; J. Forest, Museum National d'Histoire Naturelle, Paris; C. B. Goodhart, University Museum of Zool- ogy, Cambridge; H.-E. Gruner and G. Hartwich, Zoologisches Museum an der Humboldt-Universitat zu Berlin; G. Hartmann, Zoologisches Institut und Zoologisches Museum, Hamburg; R. W. Ingle, Department of Zoology, British Museum (Natural History), London; K. K. Tiwari and H. C. Ghosh, Zoological Survey of India, Calcutta; and M. Tiir- kay, Natur-Museum und Forschungsinstitut Senck- enberg, Frankfurt am Main. Part of this study, including travel to Europe in 1971 and India in 1972, was supported by the Smithsonian Institution through its Research Awards and Foreign Currency Programs. I thank Anne Cohen and Horton H. Hobbs, Jr., for their careful review of the manuscript. All of the illustrations were prepared by my wife Lilly. Family GONODACTYLIDAE Giesbrecht, 1910 Genus Gonodactylus Berthold, 1827 REMARKS.?Gonodactylus, one of the largest and most complex genera in the Stomatopoda, com- prises four distinct species groups, three of which are restricted to the Indo-West-Pacific region. There the genus is represented by the chiragra group, composed of Gonodactylus chiragra (Fabricius, 1781) and its allies, including large species (TL 90 mm or more as adults) with large ocular scales and three mid-dorsal keels or carinae on the telson (note that G. viridis Serene, 1954, redescribed be- low, may belong here in spite of its small size and ocular scales); the falcatus group, including G. fal- catus (Forskal, 1775) and similar species, compris- ing small to large species (TL 30-60 mm or more as adults) with small ocular scales and five mid- dorsal keels on the telson; and the demanii group, comprising G. demanii Henderson, 1893 and re- lated species, encompassing small species (TL usu- ally less than 50 mm as adults) with small ocular scales and the telson variously ornamented with longitudinal keels (usually three) and in some cases with spinules. In some species of the latter group the setation and shape of the uropod segments may be highly modified. It is possible that this group actually comprises species which should be assigned to two groups, one with G. demanii, and similar species, in which the telson and uropods are usually modified, and one with G. affinis and related species, in which the telson morphology more closely resembles that of members of the chiragra group. The fourth aggregate of species in the genus, the ocrstcdii group, including G. oersledii Hansen, 1895 and other American species, comprises small to large species with small ocular scales, three mid- dorsal keels on the telson, and an accessory or sup- plementary carina on the mesial side of the inter- mediate tooth of the telson. Two species of the demanii group, G. affinis de Man, 1902, and G. viridis Serene, 1954, are treated below. Gonodactylus affinis de Man, 1902 FIGURE 1 Gonodactylus chiragra var. affinis de Man, 1902:912. Gonodactylm chiragra var. confinis de Man, 1902:912, pi. 27: fig. 66. Gonodactylus chiragra var. segregatus a Lanchester, 1903:448, pi. 23: fig. 6. Gonodactylus chiragra var. segregatus b Lanchester, 1903:448, pi. 23: figs. 7, 7a. Gonodactylus chiragra var. H. affinis.?Borradaile, 1907:211 [key], 212. Gonodactylus chiragra var. K. confinis.?Borradaile, 1907:211 [key]. Gonodactylus segregatus.?Manning, 1968a:!>l, fig. 16.?Man- ning and Serene, 1968:114 [listed].?Manning, 1971:80 [key]. ?Moosa, 1973:4 [listed], 8. Gonodactylus confinis.?Manning, 1971:80 [key]. MATKRIAL.?Ternate, Molucca Islands, Indonesia; Kiiken- thal, leg.; syntypes of Gonodactylus chiragra var. affinis de Man: 1 $ . TL 21 mm; 2 9 .TL 32-34 mm (SMF 5766). Same; Kiikenthal, leg.; 1894; holotype of Gonodactylus chiragra var. confinis de Man: 1 $ , TL 22 mm (SMF). Minikoi, Laccadive Islands; reef; syntype of Gonodactylus chiragra var. segrega- NUMBER 264 tus a Lanchester: 1 $, TL 18 mm (MZC). Goidu, Goifurfe- hendu Atoll, Maldive Islands; reef; syntypes of Gonodactylus chiragra var. segregatus a Lanchester: 4 $ , TL 10-15 mm (MZC). South Male Atoll, Maldive Islands; 25 fms (46 m); syntype of Gonodactylus chiragra var. segregatus a Lanches- ter: 1 5 , TL 11 mm (MZC). Hulule, Male Atoll, Maldive Islands; reef; syntype of Gonodactylus chiragra var. segre- gatus a Lanchester: 1 juvenile, TL 8 mm (MZC). South Nilandu Atoll, Maldive Islands; 25 fms (46 m); syntype of Gonodactylus chiragra var. segregatus a Lanchester: 1 9 , TL 13 mm (MZC). South Nilandu Atoll, Maldive Islands; 19 fms (33 m); syntypes of Gonodactylus chiragra var. segregatus b Lanchester: 2,$, TL 13-20 mm; 2 $ , TL 14-15 mm; 1 juvenile, TL 9 mm (MZC). One lot containing a single female from North Male Atoll in 27-35 fms (49-67 m) could not be located in the collec- tion at Cambridge. 9 FIGURE. 1.?Gonodactylus affinis de Man, 1902: a, female syntype, TL 34 mm, Ternate, sixth abdominal somite, telson, and uropod. Gonodactylus confinis de Man, 1902, male holotype, TL 22 mm, Ternate: b, rostral plate and ocular scales; c, sixth, seventh and eighth thoracic somites, lateral view; d, sixth abdominal somite, telson, and uropod; e, submedian teeth of telson, ventral view; / , uropod, ventral view. Gonodactylus segregatus Lanchester, 1903, female syntype, TL 18 mm, Minikoi: g, sixth abdominal somite, telson, and uropod. (Setae omitted.) SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY REMARKS.?The identity of the taxa described as varieties affinis, confinis, and segregatus of Go- nodactylus chiragra could not have been deter- mined without examination of all of the types involved. Illustrations of some of the types (Figure 1) are provided here. In G. affinis the accessory median carinae of the telson are quite long and often fuse posteriorly with the median carina. The apices of the median, accessory median, and ante- rior submedian carinae each may be armed with a small spinule. de Man (1902) described these spines for G. affinis (a species that he believed to be dis- tinct from G. confinis) in which the accessory me- dian carinae are fused posteriorly with the highly inflated median carina. His G. affinis was based on four specimens, only three of which could be lo- cated, a male and two females. (The telson of one of the females is shown herein: Figure la) with relatively little inflation of the carinae of the tel- son, whereas his confinis was based on a male in which the carinae are strongly inflated (Figure Ib-f). Apparently both lots came from the same collection made at Ternate, Molucca Islands. The taxon described by Lanchester (1903) as G. chiragra var. segregatus as from several localities in the west- ern Indian Ocean is based on young specimens and females with minimal inflation of the carinae of the telson (Figure Ig). The configuration of the mid-dorsal carinae on the telson in this species superficially resembles that found in G. falcatus (Forskal) and its allies. Both de Man and Lanchester considered G. affinis to be a transitional form between "typical" G. chi- ragra (Fabricius) and "typical" G. falcatus. It is, in fact, not closely related to either but a member of a group of small species that occur in the Indo- West-Pacific region. This is a very small species; the largest individ- ual available does not exceed 34 mm in total length. Males as small as 22 mm may have well- developed secondary sexual characteristics. DISTRIBUTION.?This species is known from Indo- nesia, where it was recorded from Ternate by de Man (1902) and from several localities in the Maluku region of Indonesia in 14-32 fms (26-59 m) (Moosa, 1973); Macclesfield Bank, South China Sea (Lanchester, 1903); from Goidu, Goifurfehendu Atoll, South Male Atoll in 25 fms (46 m), Hulule, Male Atoll, North Male Atoll in 27-35 fms (49-64 m), South Nilandu Atoll in 19-25 fms (33-46 m), and Minikoi, Laccadive Islands (all Lanchester, 1903); Seychelles, Amirante Island, Saya de Malha, and Wasin, British East Africa (Wasini, Kenya) (all Borradaile, 1907); and from Madagascar in 14 m and the Comoro Islands, intertidal zone (Manning, 1968a). Gonodactylus viridis Serene, 1954 FIGURES 2a-c Gonodactylus chiragra var. viridis Serene, 1954:6, 7, 10, 74, 75, 76, fig. 13-3. fGonodactylus chiragra.?Liu, 1975:188, figs. 3-4, 3-5 [part] [not G. chiragra (Fabricius, 1781)]. MATERIAL.?KoPhuket Island, Thailand, Andaman Sea; in shale and coral, shore to 1.5 m; R. Caldwell, leg.; Aug 1973: 3 $, TL 23-41 mm; 2 9. TL 33-43 mm (USNM 150597). DESCRIPTION.?Rostral plate (Figure 2a) with width slightly greater than length or length and width subequal, apical spine relatively short; an- terolateral angles broadly rounded, acute. Ocular scales (Figure 2a) small, breadth of both less than i/3 plate width, triangular, apices angled or rounded. Sixth abdominal somite (Figure 26) with 6 carinae, narrow in juveniles, inflated in adults, each usually with apical tubercle or spinule. Telson (Figure 26) appearing broad, length and width subequal or width slightly greater, with 3 pairs of blunt mar- ginal teeth, submedians with movable apices. Sub- median, intermediate, and marginal carinae slender, unarmed dorsally. Median and anterior submedian carinae slender in juveniles and smaller females, very inflated in large males (Figure 2c), unarmed dorsally, usually unarmed posteriorly, each with posterior tubercle in smaller specimens. Accessory median carinae very short, not extending ante- riorly beyond posterior third of median carina, separate in juveniles, fusing posteriorly to form anchor in adults. Knob indistinct or absent. Row of submedian and 2 intermediate denticles present, lateral denticles absent. Uropod (Figure 26) with normal setation, proximal segment of exopod with 9-10 short, movable spines. Color completely faded in available material. MEASUREMENTS.?Males, TL 23-41 mm; females, TL 33-43 mm. Serene (1954) had more than 300 specimens available, all less than 40 mm long, in- cluding females 35 mm long with eggs. Other NUMBER 264 d FIGURE 2.?Gonodactylus viridis Serene, 1954, female, TL 33 mm, Thailand: a, ros- tral plate and ocular scales; b, sixth abdominal somite, telson, and uropod. Male, TL 41 mm, Thailand: c, sixth abdominal somite and telson. Gonodactylus chiragra (Fabricius, 1781), female, TL 64 mm, Celebes: d, rostral plate and ocular scales. (Setae omitted.) measurements of male, TL 41 mm: carapace length 11.0; rostral plate length 2.8, width 3.0; fifth ab- dominal somite width 8.5; telson length 7.2, width 7.2. REMARKS.?R. Serene named this species as a variety of G. chiragra (Fabricius) in his account of the biology of Vietnamese stomatopods published in 1954, noting that a more complete account was in preparation. It was named, figured, and distin- guished from G. chiragra, which it resembles in basic facies. Evidently it has not been recorded as a distinct taxon in more recent taxonomic papers. Apparently this species is easily recognized in the field (R. Caldwell and H. Dingle, pers. comm.), but, until the importance of the size of the ocular scales was recognized, it was extremely difficult to sepa- rate it from juveniles of G. chiragra on the basis of preserved material. It seems likely that some of the records of G. chiragra from the Indo-Malayan area should be referable to this species or to other small species of the demanii group, such as G. incipiens Lanchester, 1903 or G. micronesica Man- ning, 1971, which also have an unarmed telson and normal setation on the uropods. The G. chiragra-like telson and the small ocular scales of G. viridis will distinguish it from all other members of the demanii group. The small ocular scales serve to distinguish it from G. chiragra even in juveniles. The differences in the rostral plates and ocular scales of this species and that of G. chiragra (and the difference in ocular scale size between members of the chiragra and demanii groups) are shown in Figures 2a and 2d. Gonodactylus viridis also differs from G. chiragra in size: representatives of G. viridis are adult at a length of 35 mm, and no specimens are known to exceed 50 mm. Gonodactylus chiragra attains a length of 105 mm (Kemp, 1913), and specimens with total lengths of 70-90 mm are common. It is with some hesitation that I assign the spe- cies to the demanii group, for the basic facies and the telson of G. viridis and G. chiragra are very SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY similar. Obviously additional study of this species, based on much more material than is now available, is needed. Liu (1975) may have included G. viridis with material identified as G. chiragra in his account of some stomatopods from the Xisha Islands, China. His Figure 3 includes illustrations of the front and telson of G. chiragra (Figures 3?1, 3-2) as well as what appear to be those parts of G. viridis (Figures 3?4, 3-5). Unfortunately his account is in Chinese and I am unable to clarify his records. Serene found this species in blocks of coral in the Bay of Nhatrang, and R. Caldwell collected material in shale and coral in shallow water in Thailand; it occurred together with G. chiragra there. DISTRIBUTION.?Known with certainty from the Bay of Nhatrang, Viet Nam, and from KoPhuket Island, Thailand, in the Andaman Sea. Hoplosquilloid.es, new genus DEFINITION.?Size small, TL not exceeding 25 mm. Rostral plate (Figure 3a) sharply trispinous. Cornea (Figure 3a) elongate, not expanded later- ally. Anterior margins of lateral plates of carapace (Figure 3a) convex, extending anteriorly beyond base of rostral plate. Ischiomeral articulation of claw subterminal. Propodus of claw with proximal movable spine. Dactylus of claw unarmed. Articu- lation of propodus and dactylus inflated. Mandibu- lar palp present. Articulated anterolateral plates of abdomen (Figure 36) present. Posterior margin of sixth abdominal somite (Figure 3c) concave medially. Telson (Figure 3c,d) of unusual shape, ornamentation bizarre, dorsal and marginal arma- ture arranged in circle around almost vertical plane, anus opening in that plane above bases of marginal teeth. Posteroventral margin of telson produced into 2 primary pairs of long slender teeth, submedians with movable apices, inner and outer margins lined with long, slender, simple setae. Sub- median denticles absent, replaced by simple setae. Proximal segment of uropodal exopod (Figure 3c,/) extending beyond articulation with distal seg- ment. Movable spines on outer margin of proximal segment not markedly enlarged or strongly re- curved. Endopod slightly curved mesially. Distal segment of uropodal exopod and uropodal endo- pod (Figure 3c,/) with strong fixed spines on mesial margin. TYPE-SPECIES.?Hoplosquilloides coronatus, new species. ETYMOLOGY.?The name is derived from the Greek -oides, meaning "like," in combination with the generic name Hoplosquilla. The gender is masculine. REMARKS.?Hoplosquilloides is unique among stomatopods in having the anus situated dorsally (Figure 3c) rather than ventrally on the telson. This may well be an adaptation for burrowing, but this remains speculative, for live animals have not yet been observed. Hoplosquilloides shows closest affinities with Hoplosquilla Holthuis, 1964, represented by a single species from Ceylon, in that the uropod is armed with sharp, fixed spines. Hoplosquilloides differs from Hoplosquilla in that the mandibular palp is present and in the shape and armature of the telson, with the anus situated dorsally rather than ventrally. The telson in Hop- losquilla is typically gonodactylid, not highly modi- fied as in this new genus. Hoplosquilloides coronatus, new species FIGURE 3 MATERIAL.?St. Gilles Reef, Reunion Island; outer slope, 35 m; M. Clausade, leg., sta RE-74-22; 1974; paratype: 1 9 , TL 12 mm (SME). St. Gilles Reef, Reunion Island; outer slope, 35 m; M. Clausade, leg., sta RE-74-25; 1974; holotype: 19 , TL 18 mm (MNHNP). Same; reef flat; M. Clausade, leg., sta RE-74-55; 1974; paratype: 1 $ , TL 18 mm (USNM 156240). Grand Recif, Tulear, Madagascar; external slope, 21 m; B. Thomassin, leg., sta 230; 3 Oct 1969; paratype: 1 $ , TL 13 mm (MNHNP). Grand Recif, Tulear, Madagascar; Antseteky transect; 21-33 m; Pichon, leg., sta D-8; 2 Sep 1971; paratype: 1 $, TL 23 mm (SME). Grand Recif, Tu- lear, Madagascar; Antseteky transect, 31 m, hard bottom; Pichon leg., sta. D-13; 2 Sep 1971; paratypes: 1 $, TL 17 mm; 2 $ . TL 17-225 mm (USNM 156168). DESCRIPTION.?Ocular scales (Figure 3a) low but distinct, rounded or flattened dorsally, separate mesially. Cornea (Figure 3a) elongate, diameter greater than that of stalk. Rostral plate (Figure 3a) sharply trispinous, median spine noticeably longer than anterolaterals. Mandibular palp three-segmented. 5 epipods present. NUMBER 264 FIGURE 3.?Hoplosquilloides coronatus, new genus, new species, female paratype, TL 22.5 mm, Grand Recif: a, anterior portion of body; b, first abdominal somite, lateral view; c, sixth abdominal somite and telson; d sixth abdominal somite and telson, lateral view; e, uropod, dorsal view; /, uropod, ventral view. (Setae omitted in e, /.) Exposed thoracic and anterior 5 abdominal so- mites smooth, unarmed, lacking sharp dorsal cari- nae, abdominal somites each with marginal carina. Sixth abdominal somite (Figure 3c) with 6 longi- tudinal bosses, submedians each with posterior spinule, intermediates bluntly pointed, laterals produced into slender posterolateral spine. Ante- rior surface of lateral bosses irregular, appearing eroded. Telson (Figure 5c,d) much broader than long with short basal portion connected to long mar- ginal teeth by curved vertical surface, appearing semicircular or oval in posterior view, margin framed by short, rounded bosses dorsally and later- ally, by long marginal teeth ventrally. Each of dor- sal bosses with at least 1 posterior tubercle or spine, some bosses apically bifurcate or with secondary spinules around apical spine. Short but distinct bluntly pointed lateral tooth present, carinate lat- erally. Posterior margin with ventral 2 pairs of projections longest, submedians with movable apices, intermediates slightly shorter, with fixed apices. Submedian, intermediate, and lateral denti- cles absent. Inner and outer margins of submedian SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY and intermediate marginal teeth completely lined with single series of long, simple setae. Anus (Fig- ure 3c) visible in dorsal view, situated posterodor- sally rather than ventrally, encircled dorsally and laterally by ridge connecting inner margins of submedian teeth. Proximal segment of uropod (Figure 3e,f) with sharp, curved carina dorsally, terminating in sharp spine, 1-2 tubercles present dorsally lateral to ca- rina. Proximal segment of uropodal exopod with 11-12 movable spines laterally, distalmost slightly recurved but not enlarged, and with fixed spine ventrally (Figure 3/) overhanging articulation with distal segment. Inner margin of proximal segment of exopod (Figure 3e) smooth, lacking setae, with sharp spine at inner, distal angle. Distal segment of uropodal exopod (Figure $et,f) with normal seta- tion laterally, lacking setae but with 3-5 fixed spines mesially, and dorsally with fixed spines on short carina. Uropodal endopod (Figure 3e) with normal setation (margin evenly fringed) on outer margin, inner margin lacking setae but with strong proximal spine and several slender fixed spines distally, margin smooth between spines. COLOR.?Largely faded in preservative, but with tip of rostral plate red, anterior appendages mot- tled with dark pigment, and eyestalks brown with dark mottling. Carapace with row of dark spots across posterior third. Dactylus of claw orange or pink, less orange pigment on propodus and at merocarpal articulation. Posterior 3 thoracic so- mites each with dark spots arranged in band across somite, most prominent on sixth. All 6 abdominal somites with pair of submedian black spots, first somite also with larger, extra pair nearer to pos- terior margin, and first and fifth somites with me- dian spot. Telson with anterior pair of small, dark spots. MEASUREMENTS.?Males, TL 13-17 mm; females, TL 12-23 mm. Other measurements of female holo- type, TL 18 mm: carapace length 3.8; fifth abdomi- nal somite width 3.0; telson length 2.2, width 2.7. REMARKS.?The specimens examined are remark- ably uniform, the major variability observed being on the ornamentation of the telson. In the smaller specimens (TL 12-13 mm), the lateral projections are much less ornate than in the larger examples. There is also some minor variation in the size and position of the fixed spines on the margins of the uropod. The telson in all of the specimens appears to have a basic pattern of a median (dorsal) and seven lateral marginal projections, somewhat ob- scured by secondary ornamentation of the major projections. ETYMOLOGY.?The specific name is from the Latin corona, "crown," and -atus, "having the shape of," alluding to the shape of the telson. DISTRIBUTION.?Known only from Reunion Island and the Grand Recif, Tulear, Madagascar. It lives on coral reefs, from the reef flat to a depth of 35 m on the slope. Family LYSIOSQUILLIDAE Giesbrecht, 1910 Genus Heterosquilla Manning, 1963 Heterosquilla tricarinata (Claus, 1871) FK;URF. 4 Coronis tricarinata Claus, 1871:128, 139 [pagination from Holthuis, 1967a; pp. 21, 31 on separate]. Coronis spinosa Wood-Mason, 1875:232. Heterosquilla (Heterosquilla) tricarinata.?Holthuis, 1967a: 11 [complete synonymy].?Manning and Serene, 1968:115 [listed]. Heterosquilla (Heterosquilla) spinosa.?Holthius, 1967a: 11 [complete synonymy].?Manning and Serene, 1968:115 [listed]. MATERIAL.?Port Blair, Andaman Islands; J. Wood-Mason, don.; lectotype of Coronis spinosa: 1 $ , TL 68 mm (ZSI 3082/5). REMARKS.?In 1966 I suggested that H. spinosa from the Andaman Islands probably was distinct from H. tricarinata from New Zealand. This was based on differences between the accounts of mate- rial from the two areas, the most important being the absence of the inner spine on the basal pro- longation of the uropod in H. spinosa. Examina- tion of the lectotype of that species in 1972 revealed that the inner spine was broken on one side (Fig- ure 4e), and for this reason it was shown as an inconspicuous lobe by Kemp (1913, pi. 8: fig. 94). I can find no other differences between the speci- men from the Andaman Islands and accounts of material from New Zealand, and I believe that Coronis spinosa Wood-Mason, 1875 must be con- sidered a synonym of Coronis tricarinata Claus, 1871. The specimen figured here is the lectotype of Coronis spinosa. NUMBER 264 FIGURE 4.?Heterosquilla tricarinata (Claus, 1871), female lectotype, TL 68 mm, Andaman Islands: a, base of third walking leg, oblique ventral view; b, telson, dorsal view; c, posterior margin of telson, ventral view; d, basal part of right uropod, ventral view (basal prolongation broken); e, basal part of left uropod, ventral view (inner spine of basal prolongation broken). Keppelius, new genus DEFINITION.?Size very small, maximum length less than 30 mm. Body smooth, strongly depressed, loosely-articulated. Cornea (Figure 5a) subglobu- lar, expanded laterally beyond stalk and set slightly obliquely on it. Rostral plate (Figure 5a) subrec- tangular, anterior margin with three projections. Antennal protopod with at most 1 ventral papilla. Carapace short, narrowed anteriorly, carinae and spines absent, cervical groove scarcely indicated on lateral plates. Exposed thoracic somites lacking carinae; eighth thoracic somite lacking median ventral keel. Mandibular palp absent, 5 epipods present. Claw (Figure bb) stout, dactylus with 7 or more teeth, outer margin with basal notch; propodus stout, fully pectinate, with 4 movable spines proximally; dorsal ridge of carpus undi- vided, terminating in acute lobe; merus slender, unarmed; ischiomeral articulation terminal; is- chium three-fourths length of merus, unarmed. Endopods of pereopods two-segmented, distal seg- ment subcircular on anterior 2 legs, more ovate on third. Abdomen without longitudinal carinae or spines, articulated anterolateral plates small; sixth abdominal somite (Figure 5c) smooth dor- sally, with posterolateral spines and curved, ven- trally-directed process anterior to each uropod. Tel- son (Figure bc,d) broader than long, posterior part of surface covered with posteriorly-directed spines, marginal armature, obscured by dorsal spines, in- cluding long, movable submedian teeth and other fixed teeth and denticles. Basal segment of uropod (Figure 5c) with sharp inner and outer edges, inner terminating in distal, dorsal spine; proximal seg- ment of uropodal exopod (Figure 5r) with spatu- late spines on outer margin, rounded lobe with stiff setae on inner, distal margin; endopod (Figure 5c) oval, with strong proximal fold on outer mar- gin; inner spine of basal prolongation of uropod the longer (Figure be). TYPE-SPECIES.?Lysiosquilla hystricotelson Bar- nard, 1958. ETYMOLOGY.?It is a pleasure to name this genus for the late Keppel H. Barnard, South African Museum, Capetown. It is based on the only sto- matopod described by him. The gender is mascu- line. REMARKS.?Keppelius shares numerous features with Acanthosquilla Manning, 1963, Nannosquilla 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Manning, 1963, and Pullosquilla, new genus. It differs from Acanthosquilla in the configuration of the dorsal spines on the telson. In Acanthosquilla these spines are arranged in a transverse row across the telson surface, whereas in Keppelius there are many more spines covering a vertical shelf just above the posterior margin of the telson. Keppelius lacks distinguishable primary marginal teeth (in- termediate, lateral, and marginal) on the telson, for all of the fixed marginal teeth and denticles are subequal in size. Finally, in Keppelius the mov- able outer spines on the uropodal exopod are spatulate rather than slender. Keppelius differs from Pullosquilla in lacking an ischial spine on the claw and in having a rectangular rather than a triangular rostral plate. Keppelius agrees with Nannosquilla in basic facies, but lacks an orna- mental eave above the marginal armature of the telson and also differs in having five rather than four epipods, a dorsal spine on the antennal proto- pod, and one antennal papilla. Keppelius contains only the type-species; it is redescribed below. Keppelius hystricotelson (Barnard, 1958) FIGURE 5 Lysiosquilla hystricotelson Barnard, 1958:20, fig. 7.?Manning, 1963:319 [listed; transferred to Nannosquilla]. Nannosquilla hystricotelson.?Holthuis, 1967a:23 [synonymy]; 1967b:25, 40 [listed].?Tirmizi and Manning, 1968:5 [listed]. ?Manning and Serene, 1968:116 [listed].?Manning, 1969b:3 [list; key].?Makarov, 1971:154 [listed].?Thomassin, 1974: 303. MATERIAL.?Mombasa, Kenya; on reef exposed at S entrance to harbor; poisoned; A. J. Bruce, leg; Anton Bruun Cr. 9; 15 Nov 1964: 1 $, TL 23 mm (USNM 168779). Tulear, Madagascar; B. Thomassin, leg., sta 469: 1 $, CL 3.8 mm (MNHNP). Near Songoritelo Reef, Madagascar; B. Thomas- sin, leg., sta 444; 19 Oct 1962: 1 & , TL 19 mm; 1 9 , TL 20 mm (USNM 139073). Nosy Be, Madagascar; E beach; infra littoral, in grass; B. Thomassin, leg., sta 851; 13 Jul 1972: 1 $, TL 20.5 mm; 19 TL 23 mm (USNM 156175); and 1 $, TL 18 mm; 19 TL 24 mm (SME). Nosy Be, Madagascar; W beach; in grass; B. Thomassin, leg., sta 880; 14 Jul 1972: 19,1 carapace (MNHNP). DIAGNOSIS.?Ocular scales fused into bilobed plate. Antennal protopod with sharp antero-inter- nal spine (Figure 5a). Antennular peduncle less than half as long as carapace. Antennal scale (Fig- ure 5a) about 14 as long as carapace. Dactylus of claw (Figure 56) with 7-10 (usually 8) teeth, proximal notch on outer margin flanked by sharp proximal and obtuse distal prominences. Basal seg- ments of pereopods unarmed. Sixth abdominal somite (Figure bd) with 2 large, posteriorly directed spines on ventral surface; marginal armature of telson (Figure bc,d), on each side of midline, con- sisting of: 5-9 (usually 6-7) submedian denticles (arranged in inverted, V-shaped row), 1 movable submedian tooth, and 7 fixed lateral teeth and denticles. Proximal segment of uropodal exopod (Figure bc,e) with 3^1 blunt, movable spines on outer margin, distal 2 spatulate, and 1 stiff, spine- like seta on inner, distal angle. COLOR.?Faded in most specimens examined, but some with traces of light anterior and darker pos- terior bands on carapace and with small chromato- phores scattered over remainder of body in no particular pattern. MEASUREMENTS.?Males, TL 18-20.5 mm; fe- males, TL 20-24 mm. Barnard's type, a male, was 25 mm long. Other measurements of a female, TL 23 mm: carapace length 4.0; cornea width 0.7; ros- tral plate length 1.5, width 1.6; antennal scale length, 0.9; fifth abdominal somite width 4.0; tel- son length 2.2, width 3.4. REMARKS.?This species, with its broad rostral plate, produced anteriorly into three sharp projec- tions, and characteristically ornamented telson, can readily be distinguished from all other lysiosquil- lids. It may be confused with species of Acantho- squilla that have more than 5 spines on the dor- sal surface of the telson, but in them the telson bears fewer dorsal spines as well as a different configuration of marginal teeth. The ventral an- tennal papilla is not visible in all specimens. Thomassin (1974) found this species in sea grass beds of the fringing reef flat at Songoritelo, Mada- gascar. Like the species of Pullosquilla discussed below, K. hystricotelson is a shallow water species living on reef flats in grass beds on sand. The speci- men reported by Holthuis (1967b) came from dead coral near Massawa, Ethiopia. "DISTRIBUTION.?Western Indian Ocean, where it has been recorded from a few localities between the type-locality, Delagoa Bay, Mozambique (Bar- nard, 1958) and Massawa, Ethiopia (Holthuis, 1967b), including Songoritelo, Madagascar (Thom- assin, 1974), and the other localities reported above: Mombasa, Kenya, and Tulear and Nosy Be, Madagascar. NUMBER 264 11 FIGURE 5.?Keppelius hystricotelson (Barnard, 1958), female, TL 23 mm, Mombasa: a, anterior part of body; b, carpus, propodus, and dactylus of claw; c, sixth abdominal somite, telson, and uropod, dorsal view; d, sixth abdominal somite and telson, ventral view; e, uropod, ventral view. (Setae omitted.) Genus Neocoronida Manning, 1976 Neocoronida was established by Manning (1976a) for two species of lysiosquillids that previously had been assigned to Coronida Brooks, 1886: N. tra- churus (von Martens, 1881), from the Palau Islands and Mauritius in the Indo-West-Pacific region, and N. cocosiana (Manning, 1972), from Cocos Island in the eastern Pacific. Examination of two syntypes of N. trachurus has revealed that they are not conspecific. Neocoronida trachurus sensu stricto, from the western Indian Ocean, and N. martensi, new species, from the Palau Islands, are described and illustrated below. Key to Species of Neocoronida 1. Dorsal tubercles of telson all sharp, stellate N. trachurus (von Martens, 1881) Dorsal tubercles of telson all rounded dorsally 2 2. Rostral plate pentagonal, with slight apical projection. Ventral surface of telson tuberculate 2V. martensi, new species Rostral plate ovate, rounded anteriorly. Ventral surface of telson smooth N. cocosiana (Manning, 1972) 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Neocoronida martensi, new species FIGURE 6 Gonodactylus trachurus von Martens, 1881:93 [part]. MATERIAL.?Palau Islands, Caroline Islands, western Pacific; Piitze, leg., holotype [paralectotype of Neocoronida trachurus (von Martens, 1881)]: 1 9, TL 45 mm (ZMB 6159). DESCRIPTION.?Size small, only known specimen 45 mm long. Eye (Figure 6a) elongate, triangular, cornea noticeably broader than and set obliquely on stalk. Eye extending about to end of first segment of antennular peduncle. Ocular scales (Figure 6a) low, broad, separate. Anterior margin of ophthalmic somite produced into median spinule. Antennular peduncle about % as long as cara- pace. Antennular processes (Figure 6a) produced into blunt projections, appearing sharp in dorsal view, visible lateral to rostral plate. Shorter ramus of ventral flagellum with 21 segments in holotype. Antennal scale (Figure 6a) more than half as long as carapace. Proximal segment of antennal peduncle longer than distal segment, extending beyond eye. Protopod with blunt anterointernal projection and 1 ventral papilla. Rostral plate (Figure 6a) pentagonal, broader than long, unarmed. Lateral margins slightly con- vex, anterolateral angles broadly rounded, concave anterior margins converging on obtusely-pointed apex. Low obscure median carina present. Dactylus of claw with 4 teeth, outer margin inflated, notched basally. Dorsal ridge of carpus b FIGURE 6.?Neocoronida martensi, new species, female holotype, TL 45 mm, Palau Islands: a, anterior part of body; b, posterior part of body; c, tubercle of telson, lateral view; d, telson, ventral view; e, uropod, ventral view. (Setae omitted.) NUMBER 264 13 low, terminating in blunt projection proximal to distal margin. Mandibular palp present. 5 epipods present. Lateral process of fifth thoracic somite rounded, compressed anteroposteriorly, appearing acute in dorsal view. Fifth somite also with small ventral spinule on each side. Lateral processes of sixth and seventh thoracic somites broadly rounded. Eighth thoracic somite without noticeable median ventral keel. Endopods of perepods slender, two-segmented. Anterior 4 abdominal somites smooth dorsally, lacking tubercles or spines. Anterior portion of fifth somite smooth, remainder of surface (Figure 6b) with rounded dorsal tubercles, more numerous laterally. Fifth somite with 4 pairs of longitudinal grooves, lateralmost longest, between margin and longitudinal carina above margin; posterior margin lined with subequal tubercles or spinules, postero- lateral angles lacking spines. Sixth somite (Figure 6b) completely covered with stalked, plate-like tubercles, margins irregular, surface smooth (Figure 6c); lateral margin forming broad, carinate ridge, and with blunt posterolateral projection; posterior margin lined with spinules; acute but blunt pro- jections present ventrolaterally anterior to articula- tion of each uropod. Telson (Figure 6b,d) broader than long, posterior margin strongly convex, dorsal surface completely covered with stalked, plate-like tubercles (Figure 6c), margins irregular, surface smooth. Posterior armature of telson as illustrated. Ventral surface of telson (Figure 6d) with median patch of tubercles between anal pore and posterior margin. Uropod (Figure 6b,e) broad, flattened, anterior margin of basal segment carinate. Dorsal surface of basal segment ornamented with numerous rounded tubercles, all simple, and with short dorsal carina bearing minute apical spine. Each segment of uro- pod with dorsal tubercles, tubercles also present on ventral surface of distal segment of exopod. Outer margin of proximal segment of exopod with 9 short, movable spines, posteromesial lobe of proxi- mal segment not markedly projecting, lacking stiff setae. Distal segment of uropodal exopod longer than proximal. Endopod curved mesially, lacking strong proximal fold laterally. Spines of basal pro- longation (Figure 6e) flattened, inner longer, with 3 slender, fixed spines on inner margin. COLOR.?Completely faded in holotype. MEASUREMENTS.?Unique female holotype, TL 45 mm. Other measurements: carapace length 7.4; cornea width 2.4; antennular peduncle length 6.1; antennal scale length 4.1; rostral plate length 1.6, width 2.1; fifth abdominal somite width 9.4; telson length 5.6, width 9.1. REMARKS.?This species is known only from the holotype, which also is the paralectotype of Neoco- ronida trachurus (von Martens, 1881). Neocoronida martensi resembles iV. trachurus in most respects, differing in that the raised, plate-like tubercles on the sixth abdominal somite and telson are com- pletely smooth dorsally, and the dorsal tubercles of the fifth abdominal somite and uropods are larger, rounder, and smoother. The new species more closely resembles N. cocosiana (Manning), from Cocos Island in the Eastern Pacific, especially in the form of the plate-like tubercles, but that species has a rounded rostral plate and the ventral surface of the telson is smooth. ETYMOLOGY.?The species is named for the Ger- man carcinologist E. von Martens, who discovered N. trachurus and considered the specimen herein designated as the holotype of N. martensi as one of its members. DISTRIBUTION.?Known only from the type- locality, the Palau Islands. The depth range and habitat have not been recorded. Neocoronida trachurus (von Martens, 1881) FIGURE 7 Gonodactylus trachurus von Martens, 1881:93 [part]. Coronida trachurus.?Holthuis, 1967a:8 [references to 1967]; 1967b:40 [Red Sea references].?Makarov, 1971:153 [listed]. Coronida trachura.?Manning and Serene, 1968:115 [listed].? Tirmizi and Manning, 1968:4 [listed]. MATERIAL.?Red Sea; Jousseaume, leg.; 1898: 1 $, TL 33 mm (MNHNP). Between Grande Pass and Johny Channel, Aldabra Island; 42 m; 23 May 1953: 1 $, TL 36 mm (USNM 169780); 1 9 , TL 36 mm (MNHNP). Mauritius; G. Schnei- der, leg.; lectotype: 1 $, TL 47 mm (ZMB 6365). Mauri- tius; no other data: 2 $ (1 broken), TL 32 mm (MNHNP). Mauritius; no other data: 4 specimens, not measured (ZSI 4271/4). DESCRIPTION.?Size small to moderate, total length less than 50 mm. Eye (Figure la) elongate, triangu- lar, cornea bilobed, noticeably broader than and set obliquely on stalk. Eyes extending about to end of first segment of antennular peduncle. Ocular scales low, broad, separate. Anterior margin of 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FIGURE 7.?Neocoronida trachurus (von Martens, 1881), male lectotype, TL 47 mm, Mauritius; a, anterior part of body; b, carpus, proporius, and dactylus of claw; c, lateral processes of fifth, sixth, and seventh thoracic somites; d, posterior part of body; e, tubercle of telson, lateral view; /, telson, ventral view; g, uropod, ventral view. (Setae omitted.) ophthalmic somite produced into median spinule or tubercle. Antennular peduncle about 2/3 as long as cara- pace. Antennular processes (Figure 7a) produced into blunt projections, appearing sharp in dorsal view, visible lateral to rostral plate. Shorter ramus of antennular fiagellum with 16-19 segments. Antennal scale about half as long as carapace. Proximal segment of antennal peduncle longer than distal, extending beyond eye. Antennal pro- topod with blunt antero-internal projection and 1 ventral papilla. Rostral plate (Figure 7a) pentagonal, broader than long, unarmed. Lateral margins straight or NUMBER 264 15 slightly concave, anterolateral angles broadly rounded, concave anterior margins converging on obtusely-pointed apex. Low median carina present or absent. Dactylus of claw (Figure 7b) with 4 teeth, outer margin inflated, notched basally. Dorsal ridge of carpus low, terminating in blunt projection proxi- mal to distal margin. Mandibular palp present. 5 epipods present. Lateral process of fifth thoracic somite (Figure 7c) rounded, compressed anteroposteriorly, appear- ing acute in dorsal view, Fifth somite also with small ventral spinule or rounded tubercle on each side. Lateral processes of sixth and seventh somites (Figure 7c) broadly rounded, that of sixth somite with rounded tubercles mesial to lateral margin. Eighth somite with low, rounded median promi- nence ventrally, lacking distinct ventral keel. Endo- pods of pereopods slender, two segmented. Anterior 4 abdominal somites smooth dorsally, lacking tubercles or spines. Anteromedian portion of fifth somite (Figure 7d) smooth, remainder of surface completely covered with variously-shaped tubercles; several short, longitudinal carinae pres- ent above lateral margin, largest fusing posteriorly into marginal carina; posterior margin (Figure Id) lined with sharp tubercles or spinules, increasing in size mesially. Sixth somite (Figure Id) completely covered with stalked, stellate, or spinous tubercles (Figure 7e); lateral margin forming irregular carina terminating in blunt posterolateral projection; posterior margin (Figure Id) lined with sharp tubercles and spinules; blunt projections present ventrolaterally anterior to articulation of each uropod. Telson (Figure 7d,f) broader than long, posterior margin strongly convex, dorsal surface completely covered with stalked, stellate tubercles (Figure 7e). Posterior armature of telson as illustrated (Figure 7d,f). Ventral surface of telson (Figure 7/) with median patch of tubercles between anal pore and posterior margin. Uropod (Figure 7d,g) broad, flattened, anterior margin of basal segment carinate, dorsal surface covered with simple and stellate tubercles and with short dorsal carina terminating in blunt spine. Each segment of uropod with dorsal tubercles, tubercles also present on ventral surface of distal segment of exopod. Outer margin of proximal segment of exopod (Figure 7d) with 8-10 short, movable spines, posteromesial lobe of proximal segment not mark- edly projecting, stiff setae not discernible. Distal segment of uropodal exopod (Figure 7d) longer than proximal. Endopod (Figure 7g) curved mesi- ally, lacking strong proximal fold laterally. Spines of basal prolongation (Figure 7g) flattened, inner longer, with 1-4 slender, fixed spines on inner margin. COLOR.?Faded in most specimens, but in those from Aldabra body completely covered with nu- merous small dark chromatophores; pigment con- centrated on posterolateral angles of carapace, median area of the sixth thoracic somite, and an- terolateral plates of abdomen. Each somite with faint median and 2 lateral dark patches, body appearing striped in dorsal view. MEASUREMENTS.?Males, TL 32-47 mm; only female measured, TL 36 mm. Other measurements of male lectotype, TL 47 mm: carapace length 9.0; cornea width 2.5; antennular peduncle length ca. 6.6; rostral plate length 2.0, width 2.5; fifth abdomi- nal somite width 10.7; telson length 6.5, width 10.6. REMARKS.?Neocoronida trachurus, which appar- ently is restricted to localities in the western Indian Ocean, is readily distinguishable from the other two species of the genus by the presence of sharply stellate stalked tubercles on the sixth abdominal somite and telson; the other two species both have stalked tubercles which are rounded dorsally. Although von Martens (1881:93) mentioned two specimens from Mauritius, only one, a female 47 mm long, could be located. It is herein selected as the lectotype of Gonodactylus trachurus. The par- alectotype of G. trachurus is also the holotype of Neocoronida martensi, described above. DISTRIBUTION.?Western Indian Ocean, where it has been recorded from the Red Sea, Aldabra Island, and Mauritius. Its depth range and habitat have not been recorded. Genus Parvisquilla Manning, 1973 Parvisquilla Manning, 1973:209. DEFINITION.?Size very small, maximum length less than 20 mm. Body smooth, compact, depressed. Eye (Figure 8a) elongate, cornea faintly bilobed. Rostral plate (Figure 8a) very short, broadly rounded anteriorly. Antennular somite (Figure 8a) elongate, not overreached by rostral plate. Antennal 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY protopod lacking discernible papillae. Carapace strongly narrowed anteriorly, carinae and spines absent, cervical groove not discernible, gastric grooves very fine, poorly-marked. Exposed thoracic somites (Figure Sd) lacking longitudinal carinae; eighth thoracic somite lacking median ventral keel. Mandibular palp absent, 2 epipods present. Pro- podi of third and fourth (Figure 86) thoracic ap- pendages broader than long, lacking distinct ventral beading or ribbing. Claw (Figure 8c) stout, dactylus with 4-5 teeth, outer margin of dactylus inflated basally, with 3 proximal projections; propodus stout, proximal margin incompletely pectinate, with 3 movable spines proximally; dorsal ridge of carpus undivided, terminating in sharp spine; merus stout, unarmed; ischium much shorter than merus, unarmed. Endopods of pereopods 1-segmented, slender. Abdomen depressed but compact, smooth; articulated anterolateral plates absent; sixth ab- dominal somite (Figure Se) ornamented with irregu- lar ridges and tubercles. Telson (Figure 8 ,^/) broader than long, completely covered with raised, rounded tubercles or irregular ridges; marginal armature of telson including submedian teeth with movable apices and other fixed teeth and denticles. Uropod (Figure Se,g) stout, proximal segment with prominent outer ridge; outer margin of proximal segment of exopod completely lined with blunt spines, inner margin setose; endopod lacking prom- inent fold proximally on outer margin; basal pro- longation (Figure 8g) flattened, produced into 2 short blunt spines, inner longer. TYPE-SPECIES.?Squilla multituberculata Borra- daile, 1898. REMARKS.?This genus was established by me for two species previously assigned to Coronida, and, through an error in interpretation on my part, was transferred from the family Lysiosquillidae to the family Squillidae. The propodus of the third and fourth maxillipeds, in situ, appeared to be squillid in shape, slender and lacking the beading or ribbing characteristic of lysiosquillids. Subse- quently I dissected one of the maxillipeds of P. multituberculata to prepare the illustration pre- sented here (Figure 8b) to find that it was typically lysiosquillid in shape but lacked the ventral rib- bing typical of members of the Lysiosquillidae. Further, a better light source revealed that the gas- tric grooves, although very faint, are indeed present. Borradaile (1898, pi. 6: fig. 7), Edmondson (1921, fig. 2) and Liu (1975, pi. 1: fig. 1) all omitted the gastric grooves in their figures. Parvisquilla, which appears to be related to Coronida and Neocoronida, must be referred to the Family Lysiosquillidae. It agrees with these genera in having a stout claw with a basally-inflated dac- tylus. Its distinctive features include the greatly elongated antennular somite, the very short rostral plate, the exterior projections on the dactylus of the claw, and the suppressed mandibular palp. Two species are assigned to Parvisquilla, and one of these, P. multituberculata, is redescribed below. The other species, P. sinuosa (Edmondson, 1921), is known only from Hawaii. Key to Species of Parvisquilla Telson covered with irregular pattern of linear, curved, and scroll-like carinae P. sinuosa (Edmondson, 1921) Telson covered with erect, rounded tubercles P. multituberculata (Borradaile, 1898) Parvisquilla multituberculata (Borradaile, 1898) FIGURE 8 Squilla multituberculata Borradaile, 1898:38, pi. 6: fig. 7; 1899:403.?Manning. 1963:322 [listed]; 1973:299 [listed]. Coronida multituberculata.?Kemp, 1913:132.?Gravier, 1927: 36 [discussion].?Holthuis, 1967a:8 [synonymy].?Manning and Serene, 1968:115 [listed]. Squilla tubercitlata.?Gravier, 1927:36 [error for S. multi- tuberculata}. Parvisquilla multituberculata.?Manning, 1973:299 [discus- sion]. Parvisquilla xishaensis Liu, 1975:183, 196, pi. 1: figs. 1-6. MATERIAL?Eva Island, Tonga Islands; 21?22'S, 174?56'W; Albatross, leg.; 1899-1900: 1 $ , TL 13 mm (USNM 125330). Apia, Samoa; no other data: 2 $ , TL 11-12 mm (ZSM). Moorea, Society Islands; reef flat; M. Clausade, leg., sta MO-73-15; 1973: 1 $, TL 13 mm (SME). DESCRIPTION.?Size very small, TL of adults 13 mm or less. Body strongly depressed, surface smooth. Eye (Figure 8a) small, cornea bilobed, set obliquely on and not markedly broader than stalk. NUMBER 264 17 FIGURE 8.?Parvisquilla multituberculata (Borradaile, 1898), female, TL 13 mm, Tonga Islands: a, anterior part of body; b, propodus and dactylus of fourth maxilliped; c, claw; d, lateral processes of fifth, sixth, and seventh thoracic somites; e, sixth abdominal somite, telson, and uropod; /, margin of telson, ventral view; g, uropod, ventral view. (Setae omitted, except b.) Eyes extending almost to end of first segment of antennular peduncle. Ocular scales (Figure 8a) broad, truncate, separate mesially. Anterior margin of ophthalmic somite produced into median spinule. Antennular peduncle as long as carapace. Anten- nular processes produced into broad, blunt lobes directed almost anteriorly. Antennular somite (Figure 8a) elongate, much longer than rostral plate. Antennal protopod elongate, extending beyond antennular processes. Antennal scale very short, ovate. Proximal segment of antennal peduncle longer than distal. Rostral plate (Figure 8a) triangular, about twice as wide as long, very short, not extending to level of articulation of antennal peduncle, lacking carinae or spines. Carapace very small, strongly narrowed anteri- orly, lacking carinae and spines. Gastric grooves present, scarcely visible at magnifications of less than X 50. Anterior margins almost straight, antero- lateral angles bluntly rectangular, not projecting anteriorly. Carapace not concealing chelae, epipods, or fifth thoracic somite. Claw (Figure 8c) stout, large, almost all of ischium and merus visible in dorsal view. Dactylus sinuous, inflated basally, armed with 4 teeth; outer margin with 3 angular projections proximally, proximal largest, distalmost more obtuse than basal 2. Propo- dus deep, proximal s/4 pectinate. Dorsal ridge of carpus terminating in sharp tooth. Mandibular palp absent. 2 epipods present. Exposed thoracic somites (Figure 8d) lacking dorsal carinae or spines. Lateral process of fifth somite forming a rounded lobe, oblique to body line, compressed antero-posteriorly; blunt ventral 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY spine also present on each side. Lateral processes of sixth to eighth somites broad, truncate, flattened or rounded laterally. Pereopods stout, short, endo- pods composed of 1 segment. Abdomen smooth, broad, depressed, loosely- articulated, anterior 5 somites lacking spines or longitudinal carinae. Anterolateral plates of abdo- men absent. Fifth somite with irregular depression laterally. Sixth somite (Figure 8e) highly sculptured dorsally, with 4 pairs of longitudinal carinae, sub- medians armed, intermediates curving laterally, each terminating in blunt lobe, laterals converging with and fusing with marginals; median tubercle and 6 rounded tubercles present lateral to subme- dian carina. Posterolateral angles with rounded pro- jection. Submedian area of posterior margin lined with rounded tubercles. Telson (Figure Se,f) about 2 times broader than long, with broad, inflated, longitudinally-sulcate median boss, remainder of dorsal surface covered with large, erect, rounded tubercles. 3 pairs of sharp marginal teeth present, submedians with movable apices. Denticles slender, sharp, small, 3-5, 4-7, 1. Ventral surface smooth. Uropod (Figure Se,g) broad, basal segment with line of rounded tubercles extending to broad, rounded distal lobe. Anterolateral margin strongly convex, produced into broad, compressed, laterally carinate lobe overhanging articulation of exopod. Proximal segment of exopod shorter than distal, outer margin completely lined with 9-10 blunt spines, increasing in size distally, distal 3 slightly recurved; dorsal surface with sharp tubercles. Distal segment elongate-triangular, margin completely lined with setae. Endopod curved mesially, margin completely lined with setae, with line of 4 sharp tubercles dorsally. Basal prolongation very short, scarcely overreaching proximal segment of exopod, produced into 2 blunt spines, inner longer. MEASUREMENTS.?Only females examined, TL 11-13 mm. Borradaile (1898) reported a male and 2 females, 12-13 mm long. The female recorded by Liu (1975) was 13.6 mm long. Other measurements of female, TL 13 mm: carapace length 2.0; cornea width 0.4; antennular peduncle length 2.0; rostral plate length 0.2, width 0.4; fifth abdominal somite width 2.2; telson length 1.0, width 2.0. REMARKS.?Parvisquilla multituberailata differs from the other member of the genus, P. sinuosa, from Hawaii, primarily in having the telson orna- mented with rounded tubercles rather than irregu- lar carinae arranged in a scroll-like pattern. The few specimens of this species examined are all quite uniform, varying only in the numbers of marginal denticles on the telson. I can find no differences between the material of P. multituberculata reported herein and the species recently described from the Xisha Islands, P. xishaensis Liu, 1975. Liu's species must be con- sidered a synonym of P. multituberculata (Borra- daile). DISTRIBUTION.?Pacific Ocean, from Sandal Bay, Lifu, Loyalty Islands (Borradaile, 1898; Kemp, 1913), Shi Island, Xisha Islands, Guangdong [Kwangtung] Province, China (Liu, 1975), and now from the Tonga Islands, Samoa and Moorea, So- ciety Islands. Liu's specimen was taken on a coral reef. The depth range for the species has not been recorded. Pullosquilla, new genus DEFINITION.?Size very small, TL less than 30 mm. Body smooth, depressed, loosely-articulated. Cornea (Figure 9a) subglobular. Rostral plate (Figure 9a) triangular, unarmed anteriorly. Anten- nal protopod with ventral papilla. Carapace strongly narrowed anteriorly, carinae and spines absent, position of cervical groove indicated on lateral plates only. Exposed thoracic somites lacking longi- tudinal carinae; eighth thoracic somite lacking median ventral keel. Mandibular palp absent, 4 or 5 epipods present. Propodi of third and fourth thoracic appendages broader than long, beaded or ribbed ventrally. Claw (Figure 9b) stout, dactylus with 10 or more teeth; propodus broad, superior margin completely pectinate, with 4 movable spines proximally; dorsal ridge of carpus undivided, ter- minating in spine; ischiomeral articulation termi- nal, ischium almost as long as merus, with strong inferodistal spine on outer face. Endopods of pereopods two-segmented, distal segment ovate or subcircular on anterior 2 legs, slenderer on third. Abdomen depressed, smooth, loosely-articulated, anterolateral plates present; sixth somite unarmed dorsally; articulated anterolateral plates present; sixth somite unarmed posterolaterally, with curved, ventrally-directed process anterior to articulation of each uropod. Telson (Figure 9c, d) broader than long, smooth dorsally, dorsal surface produced into NUMBER 264 19 false eave overhanging marginal armature, obscur- ing most or all of marginal armature in dorsal view; marginal armature including row of submedian denticles, 1 movable submedian tooth, and 7 or more (?) fixed lateral teeth and denticles. Uropod (Figure 9c, e) flattened, proximal segment with dis- tal, dorsal spine; proximal segment of uropodal exopod shorter than distal, with slender, movable spines on outer margin and stiff setae on inner, dis- tal margin; endopod with strong proximal fold on outer margin; spines of basal prolongation flattened, triangular in cross-section, inner longer. TYPE-SPECIES.?Austrosquilla litoralis Michel and and Manning, 1971. ETYMOLOGY.?The name is derived from the Latin pullus, "young animal," in combination with the name Squilla, alluding to the immature ap- pearance of the species in the genus. The gender is feminine. REMARKS.?Pullosqiiilla includes three species, one of which is described as new below; all three species occur only in the Indo-West-Pacific region. A key to the species is provided below, and addi- tional records are given for P. litoralis. Pullosquilla shares the armed ischium of the claw with Austrosquilla Manning, 1966, and, because of this feature, two of the species now assigned to Pullosquilla had previously been placed in Austro- squilla. It seems likely that this feature developed independently in these genera, for in most other features Pullosquilla more closely resembles the American genus Nannosquilla Manning, 1963. These two genera share the small size as adults, the loosely-articulated body, and having the telson be- ing produced into a false eave which obscures the marginal armature. Pullosquilla differs from Nan- nosquilla in having a triangular rather than a rec- tangular rostral plate, a papilla on the antennal protopod, and a spine on the ischium of the claw. The ischium and merus of the claw are much more elongate in Pullosquilla than in Nannosquilla. Key to Species of Pullosquilla 1. Sixth adbominal somite with 2 large ventral spines. Ventral surface of telson almost com- pletely covered with spinules (Dactylus of claw with 12-15 teeth. 5 epipods present. Proxi- mal segment of uropodal exopod with 3 outer spines and 2-4 inner stiff setae) Pullosquilla thomassini, new species Sixth abdominal somite unarmed ventrally. Ventral surface of telson unarmed 2 2. Dactylus of claw with 10-13 teeth. 5 epipods present. Proximal segment of uropodal exopod with 3 outer spines and 2-5 inner stiff setae Pullosquilla litoralis (Michel and Manning, 1971) Dactylus of claw with 16-21 teeth. 4 epipods present. Proximal segment of uropodal exopod with 4 outer spines and 3 inner stiff setae Pullosquilla malayensis (Manning, 1968c) Pullosquilla litoralis (Michel and Manning, 1971) Austrosquilla litoralis Michel and Manning, 1971:237, fig. I. Austrosquilla littoralis.?Thomassin, 1974:300, 301, 307. MATERIAL.?Tiahura reef complex, Moorea, Society Islands; B. Thomassin, leg., sta TIA-3; 1973: 1$, TL 13 mm; 2 9 . TL 8.5-15 mm (SME). Data same; sta. TIA-19: 1 $ , TL 9 mm; 1 $, CL 2.5 mm (MNHNP). Data same; sta TIA-22: 1$ , TL 19 mm (MNHNP). Data same; sta TIA-23: 2$, TL 10-11 mm; 3 $ , TL 11-13 mm (USNM 169781). Aldabra Island; J. Gamble: 1 3 , TL 15 mm; 2 $ , TL 15-17 mm (BMNH). Reef, Songoritelo, Madagascar; micro-atoll flat; B. Thomassin. leg., sta 264; 26 Oct 1969: 1 9, TL 10.5 mm (SME). Grand Recif, Tulear. Madagascar; B. Thomassin, leg.; 1965: 1 ? , TL 13 mm; 3 9 ? TL 13.5-15 mm; 2 juveniles, TL 9.5 mm (1 Cl 1.8 mm) (in 6 lots; USNM 136294-136299) ( 2 g , 2 9 SME). REMARKS.?This species, originally described from a single female taken on a sand beach in the Marquesas Islands, is included here to document its occurrence off Madagascar and Aldabra Island in the western Indian Ocean, to demonstrate its abundance there and at Moorea, and to record its co-occurrence with another species of the genus, described below, at the same station on the Tiahura reef complex, Moorea. Both species occur on that reef as well as in similar habitats on the Grand Recif, Tulear, Madagascar. The specimens reported here generally agree well with the original description. There may be 10-13 teeth on the dactylus of the claw. The proximal segment of the uropodal exopod has three movable 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY spines laterally and two to five, usually three, stiff setae on the inner distal angle. Some of the smaller specimens, such as the female, TL 8.5 mm, from Tiahura, probably are postlarvae, for the eave on the telson is not yet formed and the submedian denticles of the telson are marginal and visible in dorsal view. At 10-11 mm the eave is poorly formed but visible, and it attains the adult configuration at a length of 13 mm. This species apparently prefers sandy habitats in relatively shallow water, on sand beaches or reef flats. Thomassin (1974), the first to record its oc- currence at Tulear, found it in coarse and medium sands of the holes, couloirs, and patches of the inner reef flat; he noted (p. 301) that they "reach their greatest abundance in the inner part of the moat, 250 m behind the boulder tract, generally in scattered coral-growth flats." He also found the species in the sandy bottom of a sea grass bed chan- nel opening into the lagoon at Tulear. DISTRIBUTION.?Nuka Hiva, Marquesas Islands; Tiahura, Moorea, Society Islands; Songoritelo and Tulear, Madagascar; and Aldabra Island. It occurs in shallow water on sandy bottoms. Pullosquilla thomassini, new species FIGURE 9 MATERIAL.?Tiahura reef complex, Moorea, Society Islands; B. Thomassin, leg., sta TIA-3; 1973; paratypes: 1^ , TL 10 mm (USNM 1.56276); 1 $ , TL 13 mm (SME). Tulear, Mad- agascar; sand flats; IIOE: no other data; paratype: 1$ , TL 17 mm (USNM 156254). Grand Recif, lagoon enclave, Grande Vasque (N zone), Tulear, Madagascar; 7-8 m; muddy sand; B. Thomassin, leg., sta 773; 6 Jun 1972; paratype: 1 $ , CL 2.6 mm (MN'HNP). Plage de la Batterie, Tulear, Mada- gascar; quartzy sand; B. Thomassin, leg., sta 822; 22 Jun 1972; paratvpes: 1 9 , TL 18 mm (SME); 1 juvenile, TL 11 mm. 1 abdomen (USNM 156196). Internal creek, internal slope, Grand Recif, Tulear, Madagascar; 7 m; coarse sand; B. Thomassin, leg., sta 839; 7 Jul 1972: 1 $, TL 15 mm (holotype; MNHNP); 1 $, TL 16 mm (paratype; USNM 156194). DESCRIPTION.?Cornea (Figure 9a) set slightly obliquely on stalk, overhanging stalk laterally. Eyes reaching about to end of antennular peduncle. Ocular scales fused into single, slender erect pro- jection. Antennular peduncle slightly more than 1 15 carapace length. Antennal scale (Figure 9a) about 1/4 carapace length; antennal protopod ap- parently lacking papillae. Rostral plate (Figure 9a) triangular, almost cordiform, unarmed, apex de- pressed. Dactylus of claw (Figure 96) with 12-15 teeth. 5 epipods present. Basal segments of pereo- pods each with inconspicuous posterior spine. Ven- tral surface of sixth abdominal somite (Figure 9d) with 2 strong spines on posterior margin. Telson (Figure 9c, d) broader than long, subrectangular, dorsal surface posteriorly produced into false eave (shape of eave variable, often trilobed, with small, rounded, median lobe), margin of eave with 12-22 small spines, fewer in smaller specimens; eave ob- scuring much of marginal armature in dorsal view; marginal armature consisting of transverse row of 13-17 submedian denticles, and, lateral to these, 1 movable submedian tooth and 4 curved, fixed lateral teeth, each flanked anteromesially by slender denticle; midventral surface of telson (Figure 9d) completely covered by numerous sharp fixed spines and tubercles. Basal segment of uropod (Figure 9c) with outer and dorsal carinae, dorsal carina termi- nating in 2 strong distal spines; proximal segment of uropodal exopod (Figure 9c, e) much shorter than distal, with 3 movable, spatulate spines, ar- ranged in almost transverse row on distal margin; inner distal angle of proximal segment of exopod with 2-4 (usually 3) stiff setae; distal segment of exopod and endopod very slender, curved mesially, mesial margins strongly serrate for insertion of setae; inner spine of basal prolongation of uropod longer. COLOR.?Faded in most specimens, but sometimes marked with numerous dark chromatophores scat- tered over body. In some specimens dark pigment concentrated laterally, occasionally with darker spots on sixth thoracic, fifth and sixth abdominal somites, and on telson. MEASUREMENTS.?Males, TL 10-15 mm; females, TL 13-18 mm. Other measurements of female, TL 17 mm: carapace length 3.1; cornea width 0.9; ros- tral plate length 1.3, width 1.2; antennal scale length 0.8; fifth abdominal somite width 3.2; telson length 1.7, width 3.0. REIVTARKS.?This species with its ventrally spined sixth abdominal somite, armed false eaves, and spinulose ventral surface of the telson, can be dis- tinguished easily from the other two species of the genus, P. malayensis and P. litoralis. Like the latter species, P. thomassini occurs both in Oceania and in the western Indian Ocean, and the two species have been taken together. NUMBER 264 21 d FIGURE 9.?Pullosquilla thomassini, new genus, new species, female paratype, TL 17 mm, Tulear: a, anterior part of body; b, claw; c, sixth abdominal somite, telson, and uropod; d, telson, ventral view; e, uropod, ventral view. (Setae omitted.) ETYMOLOGY.?This species is named for B. Thomassin, Station Marine d'Endoume, Marseille, who collected most of the material reported above and made it available to me for study. DISTRIBUTION.?Tiahura, Moorea, Society Islands, and localities around Tulear, Madagascar, on reef flats in grass and sand. Shallow water to a depth of 8 meters. Family SQUILLIDAE Latreille, 1803 Genus A lima Leach, 1817 Alitna orientalis, new species FIGURE 10 MATERIAL.?Takao, Formosa (Taiwan); Jun 1903; holotype: 1 $, TL 52 mm (ZSM). Mahe, Seychelles Islands; Mission Zoologique; Jul-Sep 1966; paratype: 1