Proceedings ofthe United StatesNational MuseumSMITHSONIAN INSTITUTION . WASHINGTON, B.C.Volume 124 1968 Number 3641Stoinatopod Crustacea from MadagascarBy Raymond B. ManningChairman, Department of Invertebrate Zoology
This report is based on four different collections of stomatopodsfrom the island of Madagascar (Malagasy Republic) and adjacentislands, including the Comoro Islands (Comores) and He Europa.Two of the collections were made available for study by M. MichelPichon, Centre d'Oceanographie et des Peches, Nosy Be, Madagascar,and by A. G. Humes, Boston University, at that time the field chiefon Madagascar for the International Indian Ocean Expedition. Thethird collection was made by several individuals under the auspicesof the U.S. Program in Biology, International Indian Ocean Expedi-tion; most of this series of specimens was collected by J. Rudloewhile at Madagascar on assignment from the Smithsonian InstitutionOceanographic Sorting Center. The fourth and most importantcollection was assembled by Alain Crosnier, now at the Centred'Oceanographie et des Peches, Pointe-Noire, Republic of the Congo,over a period of several years while working at the Centre d'Oceano-graphie et des Peches (O.R.S.T.O.M.) in Nosy Be, Madagascar.Together, all of these materials indicate the richness of the stomato-pod fauna of Madagascar. The collections reported herein include 28species, approximately one-half of the stomatopods known from thewestern Indian Ocean, of which six species are described as new. All10 species previously recorded from Madagascar are represented inthe collections reported in this paper. 1
2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ijmLiterature review.?Few species of stomatopods have been re-corded from the island of Madagascar. Hoffmann (1874, 1874a), inpapers on the crustaceans of Madagascar and neighboring islands,recorded the occurrence of four species from Mauritius and ReunionIslands but included no records from Madagascar. Lenz and Richters(1881) recorded two species, L. maculata (Fabricius) and G. chiragra(Fabricius), from Madagascar. Miers (1880), in the first review of thestomatopods, listed Odontodactylus scyllarus (Linnaeus) from Mada-gascar, and, in 1884, in a table of the distribution of species taken bythe Alert, included Madagascar as a locality for G. chiragra (Fabricius)but mentioned no specimens from that locality in the text. Lenz (1910)reported three species from Madagascar: Squilla nepa Latreille,Pseudosquilla ciliata (Fabricius), and Gonodactylus platysoma Wood-Mason (as G. chiragra var. acutus).Kemp (1913), in his survey of the Indo-West Pacific stomatopods,included no material from Madagascar, but he did include earlierreferences to material from there. Gravier (1920) included records forS. nepa and G. chiragra and in 1935 included records for P. ciliata,G. chiragra, and G. demanii. Also in his paper (1935) he erroneouslynoted that Lenz (1910) reported G.fimbriatus from Madagascar; Lenzonly recorded that species from Zanzibar.In an account of the stomatopods of West Africa, Monod (1925)figured the rostral plate of a specimen of L. maculata from Madagascar.In 1938 Dollfus recorded Madagascar in his paragraph on thedistribution of Gonodactylus spinosus Bigelow (as G. De Mani var.spinosus, p. 215), but no authenticated references to the occurrence ofG. spinosus in Madagascar have come to my attention. A closelyrelated species, G. lanchesteri, is recorded from Madagascar proper forthe first time in the present report.In 1941 Holthuis (p. 287) reidentilled Hoffmann's specimens of G.chiragra from Reunion as G. jalcatus (Forskal); in his section onmaterial Holthuis recorded these specimens from Reunion but men-tioned Madagascar in the text. Hoffmann mentioned only Reunion inhis account.Fourmanoir (1952, 1953) recorded G. chiragra from Madagascar and,in the second paper, gave observations on larval stages of that speciesand S. nepa. Poisson (1949), in a paper on the fauna of the Cymodoceabiotope in Madagascar, noted the occurrence of G. chiragra there.Humes (1965) recorded the occurrence of an Acanthosquilla fromNosy Be that was a host for a new cyclopoid copepod, Hemicyclopsacanthosquillae. The species of Acanthosquilla is described herein asA. humesi, new species.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 3Crosnier (1965) listed Squilla nepa and S. raphidea as occurringamong catches of penaeid shrimps off Madagascar; the latter speciesis probably Harpiosquilla harpax.In a review of the genus Odontodactylus, Manning (1967a) recordedthe occurrence of two species, 0. japonicus (de Haan) and 0. scyllarus(Linnaeus); those records were based on three specimens reportedbelow.Manning (1967b) reported Gonodactylus lanchesteri Manning fromthe Comoro Islands ; none of the seven species of Gonodactylus includedin that study were from Madagascar proper.Explanations and acknowledgments.?In general, synonymiesare shortened, including only major references and some recentpertinent references; original citations, if made by Kemp (1913), arenot repeated here. Where applicable, reference is made to Kemp's1913 monograph of the Indo-West Pacific stomatopods.In the section "Previous Kecords," earlier records of the occurrenceof each species on Madagascar are summarized. For the species notillustrated herein, a figure to aid identification of the species is citedunder the paragraph entitled "Illustration."Measurements are given in millimeters (mm). In the materialexamined, the numerals following the number of specimens denotetotal length (TL), unless carapace length (CL) is specified; the latteris given only for damaged specimens. Total length is measured alongthe midline from the apex of the rostral plate to the apices of the sub-median teeth of the telson. Carapace length is measured along themidline and does not include the rostral plate.An abdominal spine formula of "submedian, 5-6; intermediate,2-6; lateral, (1) 2-6; marginal, 1-5" indicates that the submediancarinae of the abdomen terminate in spines on the fifth and sixthsomites, the intermediate carinae terminate in spines on the secondto sixth somites, inclusive, the lateral carinae always terminate inspines on the second to somites, but these carinae occasionally arearmed on the first somite, and all five marginal carinae terminate inspines. A parenthesis is used to indicate that a carina on a certainsomite may be unarmed. A telson denticle formula of "5, 7-10, 1"indicates that on the telson margin, on each side of the midline, thereare five submedian denticles, seven to 10 intermediate denticles, andone lateral denticle. The count of teeth on the dactylus of the clawalways includes the terminal teeth.The corneal index (CI) is calculated as carapace length/cornea widthX 100.Most specimens, including all holotypes, have been deposited in theDivision of Crustacea, Smithsonian Institution (USNM); an identi-
4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124fied set of specimens also has been deposited in the Museum Nationald'Histoire Naturelle, Paris (MNHNP). The abbreviation IIOEidentifies material obtained through the International Indian OceanExpedition.I thank Michel Pichon, Arthur G. Humes, and Alain Crosnier formaking collections available for study; this report would not havebeen so complete without the large collection received from Crosnier.L. B. Holthuis, Kijksmuseum van Natuurlijke Historie, Leiden,loaned a syntype of Sqvilla harpax de Haan, and Claude Michel,Mauritius Institute, allowed me to borrow the type of Squilla juxtaora-toria Ward.The illustrations were made by my wife Lilly with the support ofthe Smithsonian Institution through its Research Awards program.Clorida Eydoux and Souleyet, 1842With the recent description of two species from India (Chhapgarand Sane, 1967) and a new species from Madagascar described herein,the genus Clorida now comprises 16 species, all of which occur in theIndo-West Pacific region. In view of the large number of species nowplaced in the genus, it is felt that the key to species presented belowwill be of some aid to other students of the group.The key does not include Sqvilla gibba Nobili, 1903, a species alignedwith Clorida latreillei Eydoux and Souleyet and its allies by previousworkers. I have transferred Nobili's species to another genus in apaper now in press (Manning, 1967c).Five species have been recorded from the Western Indian Ocean,including records given herein. Clorida latreillei has been recordedfrom southern Mozambique by Barnard (1926, 1950), and C. micro-phthalma (H. Milne-Edwards) has been recorded from Zanzibar byJurich (1904) and Stephenson (1962). Either of these two speciescould occur off Madagascar.Key to Species of Clorida
1. Mandibular palp absent 2Mandibular palp present 62. One rounded lobe present between spines of basal prolongation of uropod . 3Two rounded lobes present between spines of basal prolongation ofuropod 53. Cornea broader than stalk; rostral plate with median carina.C. incerta (Hansen, 1926)Cornea not as broad as stalk; rostral plate lacking median carina .... 44. Lateral margins of intermediate teeth of telson with prominent denticles;inner margin of basal prolongation of uropod with 3-4 spines.C. denticauda (Chhapgar and Sane, 1967)
no. 36? STOMATOPOD CRUSTACEA?MANNING 5Lateral margins of intermediate teeth of telson not denticulate; innermargin of basal prolongation of uropod with 6-9 spines.C. grant! (Stephenson, 1953)5. Sixth abdominal somite with supplementary spinules on posterior marginin addition to spines of dorsal carinae . . C. mauiana (Bigelow, 1931)Sixth abdominal somite armed at most with spines of dorsal carinae.C. fallax (Bouvier, 1914)6. First 5 abdominal somites lacking submedian carinae 7Submedian carinae present on one or more of the first 5 abdominalsomites 127. Carapace lacking anterolateral spines . . C. rotundicauda (Miers, 1880)Carapace with anterolateral spines 88. Postanal carina absent 9Postanal carina present 109. Marginal carinae of abdomen unarmed . . . C. choprai (Tweedie, 1935)Marginal carinae of at least second through fifth abdominal somites withposterior spines C. depressa (Miers, 1880)10. Cornea broader than stalk C. rniersi, new speciesCornea not as broad as stalk 1111. Width of cornea about one-third eye length; lateral margin of carapace,posterior to anterolateral spine, straight or convex; rostral plate longerthan broad C. microphthalma (H. Milne-Edwards, 1837)Width of cornea about one-half eye length; lateral margin of carapace,posterior to anterolateral spine, concave; rostral plate broader than long.C. chlorida (Brooks, 1886)12. No submedian carinae on first through third abdominal somites .... 13All 6 abdominal somites with submedian carinae 1413. Fifth and sixth abdominal somites with submedian carinae.C. merguiensis (Tiwari and Biswas, 1952)Fourth, fifth, and sixth abdominal somites with submedian carinae.C. verrucosa (Hansen, 1926)14. Lateral processes of sixth and seventh thoracic somites with posterolateralspine C. bombayensis (Chhapgar and Sane, 1967)Lateral processes of sixth and seventh thoracic somites unarmed .... 1515. Ventral surface of telson smooth on either side of postanal carina.C. latreillei Eydoux and Souleyet, 1842Ventral surface of telson tuberculate and carinate on either side of postanalcarina C. decorata Wood-Mason, 1875Clorida chlorida (Brooks, 1886)Figure 1Squilla chlorida Brooks, 1886, p. 40, pi. 2 (figs. 1-5) .?Bigelow, 1894, p. 510[key].?Kemp, 1913, p. 33.?Serene, 1952, fig. 12.Previous records.?None.Material.?1 juv., 12.1; Large Baie de Moramba, northwesterncoast of Madagascar; dredge; 30 m; muddy sand; 1 March 1958;USNM. 1 d\ 15.5; 1 9, 16.5; Banc de Pracel, western coast of Mada-gascar; dredge; 40 in; muddy sand; A. Crosnier; June 1959; USNM.2 d\ 19.5-32.0; 1 broken 9, CL 9.6; same; dredge; 65 in; sand; A.
6 PROCEEDINGS OF THE NATIONAL MUSEUMCrosnier; June 1959; MNHNP. 1 <?, 38.2; lies Mitsio, northwesterncoast of Madagascar; trawl; 64 m; muddy sand; 28 July 1958; USNM.Description.?Eye small, extending slightly beyond middle of firstsegment of antennular peduncle; stalk inflated, almost twice as longas broad; cornea small, bilobed, less than half as long as stalk; ocularscales rounded, fused along midline.
Figure 1.
?
Clorida chlorida (Brooks), male, TL 38.2, lies Mitsio: a, anterior portion ofbody; b, lateral processes of exposed thoracic somites; c, outline of lateral processes offifth and sixth thoracic somites, lateral view; d, outline of abdomen, lateral view; e, lastabdominal somite, telson, and uropod;/, uropod, ventral view. (Setae omitted).Antennular peduncle elongate, as long as carapace or nearly so;dorsal processes of antennular somite visible lateral to rostral plateas broad, sharp, anteriorly directed spines.Antennal peduncle elongate, first segment extending well beyondeye; antennal scale slender, curved, over half as long as carapace.Rostral plate broader than long, triangular, rounded anteriorly,without carinae.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 7Carapace strongly narrowed anteriorly, lateral margin concave,completely lacking median and intermediate carinae; posterior fourthof carapace usually with reflected marginal carinae only, traces oflateral carinae present or absent; posterior margin straight; antero-lateral margins slope posterolaterally to strong anterolateral spines,which do not extend to base of rostral plate.Raptorial claw stout; dactylus -with 5 teeth, outer margin of dactylusflattened; dorsal ridge of carpus with single, low, blunt spine.Mandibular palp and 4 epipods present.Exposed thoracic somites without submedian carinae, last 3 somiteswith well-formed, unarmed intermediate carinae; lateral process offifth somite a short sharp spine, directed laterally; fifth somite withblunt, tubercular, triangular or spiniform ventral projection on eachside; lateral processes of next two somites not bilobed, rounded antero-lateral^ and posterolaterally, process of sixth somite more triangularthan that of seventh; ventral keel of eighth somite a low, inconspicuousprojection.Abdomen broad, depressed, lacking submedian carinae on first 5somites; abdominal carinae spined as follows: submedian, 6; inter-mediate, 5-6; lateral, (4) 5-6; marginal, (4) 5; sixth somite with sharpspine on each side in front of articulation of uropod.Telson broader than long, with 3 pairs of sharp marginal teeth,submedians with movable apices; prelateral lobes formed dorsally, notnoticeably projecting laterally; carinae of marginal teeth short, tuber-culate dorsally; dorsal surface with short, U-shaped carinae, openposteriorly, at anterior end of line of 4 tubercles converging underposterior apex of median carina; 4 rows of tuberculate carinae presentbetween submedian and intermediate teeth, outer row anterior tocarina of intermediate tooth; single denticle present on surface anteriorto outer row, at level of U-shaped carina; small spined tubercle presentunder spine of median carina; denticles sharp, spiniform, 3, 6-7, 1;ventral surface of telson with low postanal keel.Outer margin of proximal segment of uropodal exopod with 7slender, movable spines, last extending to midlength of distal seg-ment; proximal segment of uropodal exopod with long dorsal carina;endopod slender, tapered, curved ; basal prolongation with 6-8 slenderspines on inner margin and broad, rounded lobe on outer margin oflonger inner spine.Color.?Carapace with 2 broad bands of dark pigment, gastricgrooves dark; dorsal surface of merus dark; posterior margin of cara-pace, last 3 thoracic, and all abdominal somites dark; last 3 thoracicsomites with some dark pigment on lateral processes; last 3 thoracicand first 5 abdominal somites with dorsal rectangular dark patch;pattern on telson faded, but with median triangle, apex at apical spine
8 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121
of median carina, outlined in dark pigment; uropod with dark coloron distal half of proximal segment, inner half of distal segment ofexopod, and most of endopod.Size.?Males, TL 15.5-38.2; females, TL 16.5 (largest specimenbroken); juvenile, TL 12.1. Other measurements of male, TL 38.2carapace length 7.7; cornea width 1.1 ; stalk width 1.3; eye length 2.2rostral plate length 1.3, width 1.6; fifth abdominal somite width 10.3telson length 6.2, width 8.4.Discussion.?It is with some hesitation that I assign these speci-mens to Brooks's species, one of the poorest known in the genus. Thespecimens agree in all respects with his account except that thelateral process of the fifth thoracic somite is inclined slightly forwardand there are more spines on the inner margin of the basal prolonga-tion of the uropod. Although Brooks indicated that the ventral surfaceof the telson of the type was smooth, examination of that specimenrevealed a low postanal carina that is present in the Madagascarspecimens as well.Clorida chlorida most closely resembles C. microphthalma (H. Milne-Edwards), but the latter has longer, slender eyes, a more elongaterostral plate, a broader carapace, with the lateral margins convex orstraight, and a shorter lateral process on the fifth thoracic somite.The differences in shape of the lateral margin of the carapace werenoted by R. P. Bigelow in his unpublished work on the Philippinestomatopods; it seems to be a useful character to distinguish thesetwo similar species.The young specimens are assigned to this species on the basis oftheir inflated eyes, the same number of teeth on the claw (5), theirshort rostral plate, and, in the cases of the two larger specimens, acolor pattern identical to that of all adults examined.Kemp (1913) has discounted Lanchester's (1901) record of thespecies from the Malay Peninsula, which was based on a specimenwith submedian carinae on the abdomen.Distribution.?Indo-West Pacific, from Amboina (Brooks, 1886)and off Madagascar, in depths between 27 and 64 m.Clorida fallax (Bouvier, 1914)Figure 2Squilla fallax Bouvier, 1914, p. 699; 1915, p. 308, figs. 39-42.?Barnard, 1950,p. 841 [footnote and key only].?Serene, 1954, p. G.?Stephenson and McNeill,1955, p. 241.Squilla ambigua Hansen, 1926, p. 6, pi. 1 (figs. 2 a-e).Previous records.?None.Material.? 1 broken 9, CL 5.6; Lagon de Mayotte, ComoroIslands; MYT 192 bis; A. Crosnier; August 1959; USNM.
STOMATOPOD CRUSTACEA?MANNING 9Description.?Eye small, extending beyond end of first segmentof antennular peduncle; stalk not markedly dilated, 1.8 times as longas broad; cornea broad, bilobed, broader than stalk and set trans-versely on it; ocular scales fused and sinuate along midline, acutelaterally.
wPtmv>I \
^yWtlmmm
Figure 2.
?
Clorida fallax (Bouvier), female, CL 5.6, Comoro Ids.: a, anterior portion ofbody; b, eye; c, outline of lateral processes of fifth and sixth thoracic somites; d, outlineof last three abdominal somites, lateral view; e, last abdominal somite, telson, and uropod;/, uropod, ventral view (outer spine broken). (Setae omitted).Antennular peduncle short, more than half as long as carapace;dorsal processes of antennular somite visible lateral to rostral plateas slender, anteriorly directed spines.Antennal peduncle extending to cornea; antennal scale curved,short, less than half as long as carapace.
10 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124Rostral plate longer than broad, triangular, apex rounded but acute,median carina absent.Carapace narrowed anteriorly, completely lacking median and in-termediate carinae; reflected marginal carinae present on posteriorfourth of carapace; posterior margin straight; anterolateral marginssloping laterally to short anterolateral spines.Raptorial claw slender; dactylus with 4 teeth, outer margin straight,with 2 proximal lobes, distal larger, separated by a shallow emargina-tion; dorsal ridge of carpus terminating in rounded lobe.Mandibular palp absent; 4 epipods present.Exposed thoracic somites lacking submedian carinae; last 3 thoracicsomites wTith low, unarmed intermediate carinae; lateral process offifth somite a short, slender spine, directed anterolaterally; a pah* oflong, slender ventral spines also present on fifth somite; lateralprocesses of next 2 somites subtriangular, not bilobed, rounded orangled, but unarmed posterolaterally; ventral keel of eighth somite alow, obtuse tubercle.Abdomen broad, depressed, lacking submedian carinae on first 5somites; only carinae of sixth somite armed with posterior spines;area between submedian and intermediate carinae on sixth somiteirregular; sixth somite with ventrolateral spine on each side in frontof articulation of uropod.Telson broader than long, with 3 pairs of sharp marginal teeth,subrnedians with movable apices; prelateral lobes absent; marginalcarinae and carinae of intermediate and lateral teeth smooth, carinaeof submedian teeth denticulate dorsally; entire dorsal surface of telsoncovered with carinae of varying length; dorsal submedian carinaeinterrupted posteriorly, posterior portions fusing with posterior end ofmedian carina ; dorsal surface with 1 short carina present inside carinaof submedian tooth and 4 long curved carinae, terminating in tubercles,present between submedian and intermediate teeth; anterior surfaceof telson with several other carinae of varying length, as shown infigure 2 ; median carina with distal spine overhanging smaller medianspinule; denticles 5-6, 7, 1, most sharp, outer intermediate and lateraldenticles rounded ; ventral surface with postanal keel flanked laterallyby 1 short and 1 long carina and about 14 shorter, longitudinal carinae.Outer margin of proximal segment of uropodal exopod with 5movable spines, proximal 3 sharp, distal 2 spatidate, last extendingabout to midlength of distal segment; proximal segment of exopodwith dorsal carina; endopod slender, curved; ventral surface ofprotopod with carina extending anteriorly from articulation ofendopod; inner margin of basal prolongation with 5 slender spines;inner spine of basal prolongation longer; 2 rounded lobes present
no. 3641 STOMATOPOD CRUSTACEA?MANNING 11between spines of basal prolongation, inner larger, apex deflecteddorsally.Color.?Posterolateral angles of carapace and lateral portions ofbody black; telson with lateral black patches; nropodal exopod withdark spot at articulation of distal segment.Size.?Only specimen examined, a female, broken. Measurements:carapace length 5.6; cornea width 1.3; eye length 1.8; stalk width 1.0;rostral plate length 1.2, width 0.8; fifth abdominal somite width 6.5;telson length 3.8, width 5.4.Discussion.?The present broken specimen agrees very well withthe limited accounts of S. fallax given by Bouvier (1914, 1915) andthe detailed account of S. ambigua given by Hansen (1926). Theiraccounts differ only in the extent of carination of the telson and thearmature of the marginal carinae of the abdomen. Bouvier's figure(1915) indicates that most of the dorsal carinae of the telson arebroken, whereas the carinae on the telson of Hansen's specimen aremostly entire. In the broken specimen from the Comoro Islands theyare divided proximally, entire distally. On another specimen in theU.S. National Museum, from the Solomon Islands, the carinae of thetelson are broken mostly into tubercles. In view of the normal var-iation in telson ornamentation in this genus, the extremes exhibitedby Hansen's specimen and Bouvier's as well must be consideredwithin the expected range of variation.Although the specimen from the Comoro Islands lacks marginalspines on the fifth abdominal somite, as in S. fallax, the larger speci-men from the Solomon Islands has this spine, which was also presentin Hansen's type.Finally, the dark lateral portions of the body, mentioned by bothHansen and Bouvier, are present in the specimens from both theIndian Ocean and Pacific Ocean.Distribution.?Indo-West Pacific, from Mauritius (Bouvier, 1914,1915), Madagascar, Dangar Besaar, Indo-Malaya (Hansen, 1926),Viet Nam (Serene 1954), New South Wales, Australia (Stephensonand McNeill, 1955), and the Solomon Islands.Clorida miersi, new speciesFigure 3Holotype.?1 d\ 32.8; Banc de Pracel, western coast of Mada-gascar; 65 m; sand; A. Crosnier; Jane 1959; USNM 124091.Description.?Eye small, extending to end of first segment ofantennular peduncle; stalk slightly inflated, not as broad as cornea;cornea small, bilobed, breadth about two-thirds length of eye; ocularscales rounded, fused along midline.
12 PROCEEDINGS OF THE NATIONAL MUSEUM vol. imAntennular peduncle elongate, over half as long as carapace; dorsalprocesses of antennular somite visible lateral to rostral plate as slender,sharp, anteriorly directed spines.Antennal peduncle not extending beyond eye; antennal scale slender,curved, less than half as long as carapace.Rostral plate longer than broad, margins proximally subparallel,distally convergent on rounded apex; median carina absent.Carapace strongly narrowed anteriorly, completely lacking medianand intermediate carinae; posterior fourth of carapace with reflectedmarginal carinae only; posterior margin straight; anterior marginsslope posterolaterally to strong anterolateral spines, which do notextend to base of rostral plate.Raptorial claw slender: dactylus with 5 teeth, outer margin flat-tened, with proximal notch flanked distally by rounded lobe; dorsalridge of carpus terminating in low, rounded tubercle.Mandibular palp and 4 epipods present.Exposed thoracic somites lacking submedian carinae, least 3 somiteswith well-formed, unarmed intermediate carinae; lateral process offifth somite a short, sharp, slender, anterolaterally directed spine;fifth somite with ventral tubercle under each lateral spine; lateralprocesses of sixth and seventh somites not bilobed, obliquely truncate,rounded anterolaterally and posterolaterally; ventral keel of eighthsomite prominent, rounded, projecting ventrally.Abdomen broad, depressed, first 5 somites lacking submediancarinae; abdominal carinae spined as follows: submedian, 6; inter-mediate, 5-6; lateral, 6; marginal, 5; sixth somite with ventrolateralspine in front of articulation of each uropod.Telson broader than long, with 3 pairs of sharp marginal teeth,submedians with movable apices; prelateral lobes absent; carinae ofsubmedian and intermediate teeth nodulose, marginal carinae smooth;telson ornamented dorsally with a series of anterior tubercles, a sub-median line of tubercles on each side of median carina convergingunder its apex, and 6-7 short, curved lines of tubercles; mediancarina with slender spine and ventral tubercle under spine; denticlessharp, 2-3, 7-8, 1; ventral surface of telson with postanal keel.Outer margin of proximal segment of uropodal exopod with 6slender, curved spines, last extending about to midlength of distalsegment; proximal segment of exopod with longitudinal dorsal carina;endopod slender, curved; basal prolongation with 7 slender spines oninner margin and broad rounded lobe on outer margin of longer innerspine.Color.?Almost completely faded; some dark chromatophores pres-ent on anterior appendages, arranged in lines on propodus of claw;
STOMATOPOD CRUSTACEA?MANNING 13
uropod with traces of dark pigment on inner half of exopod and distalhalf of endopod.Size.?Male holotype, only specimen examined, TL 32.8. Othermeasurements: carapace length 5.2; cornea width 1.3; eye length 1.8;
Figure 3.
?
Clorida miersi, new species, male holotype, TL 32.8, Banc de Pracel: a, anteriorportion of body; b, eye; c, outline of lateral processes of fifth, sixth, and seventh thoracicsomites; d, outline of fifth thoracic somite, lateral view; e, outline of abdomen, lateralview; /, last abdominal somite, telson, and uropod; g, basal prolongation of uropod,ventral view. (Setae omitted).
14 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124
stalk width 1.1; rostral plate length 1.4, width 0.9; fifth abdominalsomite width 7.3; telson length 3.9, width 5.3.Discussion.-?Cloridamiersi closely resembles C. verrucosa (Hansen)in most features but shows several important differences, as follows:(a) the eyes are of different shape, the stalk is not as long and is lessexpanded laterally, and the broader cornea is set transversely on thestalk; (b) there are no submedian carinae on the first five abdominalsomites, but they are present on the fourth and fifth somites in C.verrucosa; (c) fewer abdominal carinae are armed (in C. verrucosa theabdominal carinae of the fourth, fifth, and sixth somites are all pro-vided with spines) ; and (d) the anterior lobe of the lateral process ofthe fifth thoracic somite is spiniform in C. verrucosa, rounded in C.miersi.The large eyes and elongate rostral plate will immediately dis-tinguish this species from the others known from the western IndianOcean, C. clorida (Brooks), C. fallax (Bouvier), C. latreillei Eydouxand Souleyet, and C. microphthalma (H. Milne-Edwards).Etymology.?The species is named for E. J. Miers, the Britishcarcinologist who made valuable early contributions to our knowledgeof the stomatopods.Distribution.?Known only from the type-locality, off the westcoast of Madagascar.Harpiosquilla Ilolthuis, 1964Four species, all from the Indo-West Pacific region, have beenassigned to this genus; all four occur in the western Indian Ocean.Balss (1910) reported H. raphidea from off East Africa and Chopra(1939) recorded H. annandalei from the Gulf of Oman. Two species,H. harpax and one described as new, are reported herein. The followingkey will serve to distinguish all of the species.Key to Species of Harpiosquilla
1. Fifth thoracic somite with a lateral spine 2Fifth thoracic somite rounded laterally 32. Submedian carinae of fifth abdominal somite armed posteriorly ; distal segmentof uropodal exopod black with a white midrib . H. annandalei (Kemp, 1911)Submedian carinae of fifth abdominal somite unarmed; distal segment ofuropodal exopod with inner half only dark, not black.H. raphidea (Fabricius, 1798)3. Carapace with median carina; first 5 abdominal somites with submediancarinae H. harpax (de Haan, 1844)Carapace lacking median carina; first 5 abdominal somites lacking submediancarinae H. melanoura, new species
no. 3641 STOMATOPOD CRUSTACEA?MANNING 15Harpiosquilla harpax (de Haan, 1884)Figure 4Squilla harpax de Haan, 1844, pi. 51 (fig. 1).?Tiwari and Biswas, 1952, p. 358,figs. 3b, d, f?Barnard, 1955, p. 49.?Ingle, 1963, p. 18, figs. 9, 59.?Holthuis,1964, p. 140 [description of Harpiosquilla].Squilla raphidea.?Crosnier, 1965, p. 61 [listed].Not Harpiosquilla harpax.?Manning, 1966, p. 87, fig. 1 [an undescribed species],Harpiosquilla harpax.?Manning, 1967, p. 103.Previous records.?Crosnier, 1965.Material.? 1 9, 118.2; Ambaro Bay, Nosy Be; trawl; 2-5 m; M.Pichon; 8 September 1964; USNM. Id1 , 171.0; Baie de Narendry,northwestern coast of Madagascar; trawl; 6 m; mud; A. Crosnier;February 1958; USNM. 1 9, 141.5; same; MNHNP.Description.?-Eye large, T-shaped, cornea bilobed, set almosttransversely on stalk; eyes not extending past end of first segment ofantennular peduncle; ocular scales truncate, situated laterally; an-terior margin of ophthalmic somite rounded; corneal indices 269-358.Antennular peduncle slender, as long as or slightly shorter thancarapace; dorsal processes of antennular somite slender, tapered,directed anterolaterally.Rostral plate triangular, longer than broad, without carinae;lateral margins sinuous, convex posteriorly, convex anteriorly, con-verging on short, blunt apex that extends just beyond edge of an-tennular somite.Carapace narrowed anteriorly, anterior width less than one-halfmedian length; anterolateral spines strong but not extending to baseof rostral plate; median carina present, not bifurcate at either end;intermediate carinae not extending to anterior margin.Mandibular palp and 5 epipods present.Raptorial claw large, propodus almost half again as long as cara-pace; dactylus with outer margin evenly curved in female, withproximal obtuse angle in male, inner margin with 8 teeth; propoduswith erect spines and tubercles on margin opposing dactylus; dorsalridge of carpus undivided.Exposed thoracic somites with, at most, vestiges of submediancarinae, last three somites with prominent unarmed intermediatecarinae; lateral process of fifth somite rounded, obscure, a ventralspine, with convex posterior lobe, present on each side; lateral processof sixth somite bilobed, anterior lobe low, rounded, posterior tri-angular, with spiniform apex; lateral process of seventh somite withirregular margin, at most an anterior tubercle present, postero-lateral^ spiniform; ventral keel on eighth somite rounded, inclinedposteriorly.
16 PROCEEDINGS OF THE NATIONAL MUSEUMAbdomen with submedian carinae of first 5 somites low, poorlydeveloped, but present; remainder of abdominal carinae well-formed,spined as follows: submedian, 6; intermediate, 2-6; lateral, 1-6;marginal, 1-5; sixth somite with sharp spine in front of articulationof each uropod and with median ventral carina.
Figure i.-Harpiosguilla harpax (de Haan), female, TL 118.2, Ambaro Bay: a, anteriorportion of body; b, outline of eye; c, outline of rostral plate; d, outline of lateral processesof exposed thoracic somites; e, last two abdominal somites, telson, and uropod;/, basalprolongation of uropod, ventral view. (Setae omitted).
no. 3641 STOMATOPOD CRUSTACEA?MANNING 17Telson as broad as long, appearing elongate; carinae of marginalteeth short, rounded, slightly nodulose dorsally; 3 pairs of marginalteeth present, prelateral lobe formed dorsally but not projectinglaterally; marginal carina over twice as long as carina of lateral tooth;denticles subequal, sharp, 4-7, 10-12, 1 ; ventral surface with postanalkeel extending about halfway between anus and posterior margin.Uropod with 9 movable teeth on outer margin of proximal segmentof exopod, last short; lobe on outer margin of inner spine of basalprolongation prominent, rounded, margin concave.Color.?Anterior edge of antennal scale dark; carinae, grooves,and posterior margin of carapace dark; thoracic and abdominalsomites with anterior and posterior dark, transverse lines, posteriorbroader and darker, and intermediate and marginal carinae outlinedwith dark pigment; telson with pair of anterior, submedian, roundblack spots; carinae and margins of uropod with dark chromatophores,particularly inner margin of exopod; endopod with dark pigmentlining both edges of distal half, central portion clear.Size.?Male, TL 171.0; females, TL 118.2-141.5. Other measure-ments of male, TL 171.0: carapace length 32.2; cornea width 10.4;rostral plate length 5.6, width 5.1 ; fifth abdominal somite width 32.8;telson length 30.4, width 30.4.Discussion.?The presence of a median carina on the carapace,submedian carinae on the abdomen, and the lighter colored uropodsdistinguish this species from H. melanoura, new species, describedbelow. Other differences are discussed under the account of the newspecies.Harpiosquilla harpax lacks the lateral spine on the fifth thoracicsomite, which is characteristic of H. raphidea (Fabricius), and lacksthe armed intermediate carinae of the thoracic somites, which char-acterize H. annandalei (Kemp) and which may occur also in verylarge specimens of H. raphidea. In their account separating H. harpaxfrom H. raphidea, Tiwari and Biswas (1952) also pointed out otherdifferences between the two species.The specimen from Australia assigned to this species by Manning(1966) belongs to an undescribed species, an account of which is inpreparation.Through the kindness of L. B. Holthuis, Rijksmuseum van Natuur-lijke Historie, Leiden, I Avas able to study one of the syntypes ofSquilla harpax de Haan and compare it directly with the specimens ofH. harpax and H. melanoura, new species, reported here. The specimen,a male, TL 152.5, from Japan, RMNH 28H, is here selected as thelectotype of Squilla harpax. It agrees in all details with the specimensof H. harpax from Madagascar.273-102?67 2
18 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124The corneal indices of the two females of H. harpax reported here,CL 21.0 and 28.5, are 269 and 320; in the two males (including thelectotype), CL 29.1-32.2, they are 316 and 358, respectively. Tiwariand Biswas (1952) reported a range of 2.7-3.2 (270-320) in theirmaterial. Not enough material is available to determine if there areany sexual or geographic differences in the corneal indices of thisspecies.The eyes of H. melanoura, new species, appear to be larger, and therange of corneal indices, 229-252, indicates that the cornea is appre-ciably broader in that species.The corneal index of the specimen reported from Australia byManning (1966), a female, CL 30.6, was 336, which falls within therange observed for H. harpax. In the Australian specimen, however,the eye appears to be more inflated than in H. harpax.Distribution.?Indo-West Pacific, from the Red Sea and EastAfrica to Japan. Harpiosquilla melanoura, new speciesFigure 5Holotype.? 1 cf, 128.6; Banc de Pracel, western coast of Mada-gascar; 55 m; sand; A. Crosnier; July 1959; USNM 124092.Paratypes.? 1 9, 135.6; Banc de Pracel, western coast of Mada-gascar; 55 m; muddy sand; A. Crosnier; June 1959; USNM 124093.1 &, 101.6; 1 9, 126.1; same; MNHNP.Description.?Eye large, T-shaped, cornea bilobed, set almosttransversely on stalk; eyes extending anteriorly almost to end offirst segment of antennular peduncle; ocular scales obliquely truncateor rounded, situated laterally; anterior margin of ophthalmic somiterounded; corneal indices 229-252.Antennular peduncle slender, slightly longer than carapace; dorsalprocesses of antennular somite broad, each tapered to an acute apex,directed anterolaterally.Antennal scale slender, curved, two-thirds as long as carapace.Rostral plate triangular, longer than broad, without carinae; lateralmargins sinuous, convex posteriorly, concave anteriorly, convergingon slender, blunt apex; plate extending just beyond dorsal margin ofantennular somite.Carapace narrowed anteriorly, anterior width less than one-halfmedian length; anterolateral spines strong but not extending to baseof rostral plate; median carina completely absent; intermediatecarinae not extending to anterior margin.Mandibular palp and 5 epipods present.Raptorial claw large, propodus slightly less than half again as long ascarapace; dactylus with 8 teeth, outer margin of dactylus evenly
NO. 3641 STOMATOPOD CRUSTACEA?MANNING 19
curved in female, with proximal, obtuse, angular prominence inmale; propodus with series of upright spines on upper margin; dorsalridge of carpus undivided.
Figure S.?Harpiosquilla melanoura, new species, male holotype, TL 128.6, Banc de Pracel:a, anterior portion of body; b, eye; c, lateral processes of exposed thoracic somites; d, lasttwo abdominal somites, telson, and uropod; e, basal prolongation of uropod, ventralview. (Setae omitted).Exposed thoracic somites lacking submedian carinae, last 3 somiteswith short, unarmed intermediate carinae; lateral process of fifththoracic somite obscure, unarmed; fifth somite with slender ventralspine on each side, margins sinuous, apex very sharp; lateral process
20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121
of sixth somite bilobed, anterior lobe obtusely rounded, posteriortriangular, apex spiniform; lateral process of seventh somite not con-spicuously bilobed, margin sinuous, triangular posterolateral apexspiniform; ventral keel of eighth somite inclined posteriorly, roundedanteriorly, angled or rounded posteriorly.Abdomen lacking submedian carinae on first 5 somites; remainderof abdominal carinae well-formed, spined as follows: submedian, 6;intermediate, 2-6 ; lateral, 1-6 ; marginal, 1-5 ; sixth somite with sharpposterolateral spine in front of articulation of uropod and short,median, ventral keel.Telson noticeably longer than broad, median carina thick, termi-nating in slender posterior spine; carinae of marginal teeth short,rounded, slightly irregular dorsally; 3 pairs of marginal teeth present,prelateral lobes formed dorsally but not projecting laterally; marginalcarina almost 3 times as long as carina of lateral tooth; denticlessubequal, angular but not sharp, 5, 9-10, 1; ventral surface withshort postanal keel, not extending halfway from anus to posteriormargin.Uropod with 9 graded, movable spines on outer margin of proximalsegment of exopod, last short; lobe on outer margin of inner spine ofbasal prolongation prominent, rounded, inner margin concave.Color.?Anterolateral angle of antennal scale black; carapace withU-shaped median dark patch, open anteriorly, anteror to cervicalgroove; posterior margin of carapace, last 3 thoracic somites, and first5 abdominal somites black; telson with pair of proximal, submedianblack squares, convergent rows of pits on surface black; distal seg-ment of uropodal exopod black, with small, clear posterolateral area;distal two-thirds of uropodal endopod black.Size.?Males, TL 101.6-128.6; females, TL 126.1-135.6. Othermeasurements of male holotype: carapace length 24.4; cornea width10.2; rostral plate length 4.6, width 4.1; fifth abdominal somite width24.6; telson length 24.5, width 21.7.Discussion.?H. melanoura most closely resembles TI. harpax (deHaan), but it differs in several important features, as follows: (a) theeyes are noticeably larger; (b) the median carina of the carapace isabsent; (c) submedian carinae are absent on the first five abdominalsomites; (d) the telson is slenderer and longer; (e) the postanal keel isshorter, less than half the distance between the anus and the posteriormargin of the telson; and (f) the distal portions of the uropod areblack. Specimens of the two species can be separated immediately bythe differences in color of the uropod alone.Harpiosquilla raphidea (Fabricius) is a larger species, in which thelateral process of the fifth thoracic somite is spined; it is unarmed inH. melanoura. The only other known species of Harpiosquilla, H.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 21
annandalei (Kemp), differs from H. melanoura in having the inter-mediate carinae of the last three thoracic somites armed with aposterior spine.The four specimens of H. melanoura described here have been com-pared with the leetotype of H. harpax (de Haan).Etymology.?The name is derived from the Greek, 'melanos,"meaning "dark," and from "oura," meaning "tail," referring to thedark uropods of the species.Distribution.?Known only from the type-locality, Banc dePracel, off the west coast of Madagascar; all specimens were takenat 55 m. Squilla Fabricius, 1787Ten species of Squilla have been recorded in the western IndianOcean from localities between the Red Sea and southern Mocambique,and five of these occur off Madagascar. The five species not nowrecorded from Madagascar are: Squilla investigatoris Lloyd, 1907,from the Gulf of Aden (Chopra, 1939; Ingle, 1963); S. mikado Kempand Chopra, 1921, from Zanzibar and Mocambique (Chopra, 1939;Barnard, 1950; Manning, 1965); S. massavensis Kossmann, 1880,from the Red Sea and Zanzibar(?) (Ingle, 1963) ; S. minor Jurich,1904, from Zanzibar; and S. woodmasoni Kemp, 1911, from Zanzi-bar^) and Mozambique (Kemp, 1913; Barnard, 1962). With theexception of S. massavensis, which may not occur south of the RedSea (see discussion below), any of these species could occur offMadagascar. Squilla carinata Serene, 1950Figure 6Squilla multicarinata.?Dollfus, 1938, p. 196, fig. 1.? Gravier, 1938, p. 174, fig. 4[not S. multicarinata White].Squilla carinata Serene, 1950, p. 571; 1954, pp. 6, 8.?Ingle, 1963, p. 17, figs.6-8, 10-12, 67.Previous records.?None.Material.? 1 9, 65.1 ; Banc de Pracel, western coast of Madagascar;55 m; muddy sand; A. Crosnier; June 1959; MNHNP. 1 d% 88.4;Baie d'Ambaro, Madagascar; 26 m; shelly sand; A. Crosnier; June1959; USNM.Description.?Body covered with short, longitudinal carinae;antennular peduncle longer than carapace and rostral plate combined;eye of moderate size, cornea set obliquely on stalk, stalk with severalirregularly placed dorsal carinae; ocular scales bifurcate, apicesrounded or acute; rostral plate appearing elongate, with long mediancarina and carinate lateral margins; carapace narrowed anteriorly,covered with longitudinal carinae, median carina with anterior
22 PROCEEDINGS OF THE NATIONAL MUSEUMbifurcation, posterior margin produced into median point; antero-lateral spines of carapace strong but not extending to base of rostralplate; mandibular palp and 4 epipods present; raptorial claw slender,dactylus with 6 teeth, outer margin of dactylus flattened but notconspicuously sinuate; dorsal ridge of carpus undivided; fifth thoracicsomite irregularly carinate, with longitudinal and transverse carinae;anterior portion of lateral process of fifth somite directed antero-laterally, apex blunt; posterior lobe slender, apex rounded; submediancarinae of last 3 thoracic somites armed posteriorly, at least 4 of carinaebetween submedians also armed ; lateral process of sixth somite bilobed,anterior lobe large, apex obliquely truncate, posterior lobe triangular,
Figure 6.
?
Squilla carinata Serene, male TL88.4, AmbaroBay: anterior portion of body.
slenderer; lateral process of seventh thoracic somite bilobed, bothlobes triangular, anterior lobe smaller; ventral keel on eighth thoracicsomite low, triangular; each abdominal somite with numerous longi-tudinal carinae, last somite with fewest; abdomen with submedian,intermediate, and lateral carinae spined posteriorly on all 6 somites,marginals of first 5 somites also armed; on first 5 somites median andup to 4 other carinae between submedians also armed; on fifth somiteonly, 1-2 carinae on each side between intermediate and lateralcarinae spined; remainer of abdominal carinae unarmed; telson slender,longer than broad, with 4 pairs of sharp marginal teeth and 12 ormore carinae of varying length on either side of median carina;
no. 3G4i STOMATOPOD CRUSTACEA?MANNING 23denticles rounded, 4-5, 8-9, 1; ventral surface of telson with postanalkeel flanked by numerous longitudinal carinae; uropod slender,elongate, all segments longitudinally carinate; penultimate segmentof exopod with 11 graded, movable spines, last not extending to mid-length of distal segment; basal prolongation of uropod with series ofshort spines on inner margin, lobe on outer margin of inner spinesmall, acute.Color.?Posterior margin of carapace, last 3 thoracic and first 5abdominal somites lined with black; body carinae with faint traces oflongitudinal lines of dark pigment; telson with bases of intermediate,lateral, and marginal teeth dark, dark pigment on telson terminatinganteriorly at a line across level of posterior spine of median carina;uropodal exopod with distal fourth of proximal segment and most ofdistal segment black, apex clear; distal third of uropodal endopodblack.Size.?Male, TL 88.4; female, TL 65.1. Other measurements ofmale: carapace length 19.8; cornea width 4.4; rostral plate length 3.0,width 2.6; telson length 17.2, width 14.8.Discussion.?The description given above is based on the twospecimens reported herein. Judging from other accounts in the litera-ture, this species exhibits some variation in ornamentation and sina-tion of the body carinae. Dollfus (1938, fig. 7) showed that in onespecimen from the Gulf of Suez several of the body carinae, otherthan those reported here, terminated in spines.The present material agrees with Serene's account in having carinaeon the eyestalks, bifurcate ocular scales, and a mandibular palp.Serene did not go into any detail on the other features of his materialbut did mention that the first two of these characters would differenti-ate this species from others. The Madagascar specimens agree wellwith the accounts of the species given by Gravier (1938), Dollfus(1938), and Ingle (1963).Distribution.?Indo-West Pacific, from the Gulf of Suez, Mada-gascar, and Viet Nam; the species has not been recorded previouslyfrom Madagascar. Squilla gonypetes Kemp, 1911Figure 7Squilla gonypetes.?Kemp, 1913, p. 54, pi. 4 (figs. 42-44).?Ingle, 1963, p. 15,figs. 1, 5, 14.?Manning, 1965, p. 250, pi. xi (fig. b) [older references].Previous records.?None.Material.? 1 cf, 53.1; 1 9, 45.2; Banc de Pracel, western coast ofMadagascar; 55 m; muddy sand; A. Crosnier; June 1959; USNM.1 d\ 51.4; 1 9, 38.8; same; MNHNP. 1 cf postlarva, 14.8; Large Baie
24 PROCEEDINGS OF THE NATIONAL MUSEUMMoramba, northwestern coast of Madagascar; dredge; 30 m; muddysand; A. Crosnier; 1 March 1958; USNM.Diagnosis.?Eye small, cornea set obliquely on stalk; rostral platewithout median carina; anterior bifurcation of median carina ofcarapace obscure, interrupted in some specimens; dactyl us of raptorialclaw with 5 teeth; dorsal ridge of carpus of claw undivided; inferodistal
Figure 7.
?
Squilla gonypetes Kemp, male, TL 53.1, Banc de Pracel: a, anterior portion ofbody; b, outline of lateral processes of fifth, sixth and seventh thoracic somites.
angle of merus of claw unarmed; mandibular palp and 4 epipods pres-ent; lateral processes of fifth to seventh thoracic somites bilobed,anterior lobe of process of seventh somite well developed; abdominalcarinae spined as follows: submedian, 5-6; intermediate, 4-6; lateral,3-6; marginal, 1-5; telson with 4, 7, 1 denticles, outer submedian andouter intermediate larger than remainder; ventral surface of telsonwith short postanal keel.Color.?Thoracic and abdominal somites with dark, irregularposterior line; second abdominal somite with dark transverse patch,fifth somite with pan of dark, triangular submedian patches; sixthabdominal somite black posterolaterally; telson with rectangularanterior and triangular posterior dark patches along median carina;
no. sen STOMATOPOD CRUSTACEA?MANNING 25
uropodal exopod with distal portion of proximal segment and innerhalf of distal segment with scattered dark chromatophores ; distalhalf of endopod dark.Discussion.?-These specimens agree in most details with Kemp's(1913) account of the species, but differ in the following features:(a) the rostral plate appears to be more elongate; (b) the branches ofthe anterior bifurcation of the median carina of the carapace arepresent although obscure; (c) the dactylus of the claw is curved morestrongly; (d) the posterior lobe of the lateral process of the sevenththoracic somite is not acute but rounded; (e) the intermediate carinaeof the third abdominal somite and the lateral carinae of the secondsomite are unarmed; and (f) the submedian dark squares of the fifthabdominal somite are represented in these specimens by triangles.Kemp noted that one specimen from Madras differed in these respectsfrom other material available to him. This species still is known toopoorly to determine whether or not these differences reflect normalvariation, or whether or not two or more species are present.Distribution.?Indo-West Pacific, from scattered localities be-tween the Red Sea and Japan.Squilla hesperia, new speciesFigure 8ISquilla nepa.?Miers, 1880, p. 25 [part; specimen from Zanzibar].tSquilla woodmasoni.?Kemp, 1913, p. 74 [part; specimens from Zanzibar re-ported by Miers (1880), reidentified].
'tSquilla massavensis.?Ingle, 1963, p. 15 [part; specimens from Zanzibar reportedby Miers (1880) and Kemp (1913), reidentified].Holotype.? 1 9, 116.6; Baie de Tsimipaika, northwestern coast ofMadagascar; trawl; 15 m; muddy sand; A. Crosnier; March 1959;USNM 124094.Description.?Eye large, cornea bilobed, set very obliquely onstalk; prominent lateral projection present on each stalk; eyes ex-tending to end of first segment of antennular peduncle; anteriormargin of ophthalmic somite obtusely rounded; corneal index 364.Antennular peduncle slightly shorter than carapace; dorsal pro-cesses of antennular somite visible lateral to rostral plate as slender,acute projections, directed anterolaterally.Antennal scale slender, curved, over one-half as long as carapace.Rostral plate subquadrate, as long as broad, margins sloping t<>rounded apex; neither median carina nor median tubercle present;lateral margins carinate.Carapace broad, anterior width more than one-half median length,excluding rostral plate; anterolateral spines of carapace not extendingto base of rostral plate; anterior portion of median carina of carapace
26 PROCEEDINGS OF THE NATIONAL MUSEUMlacking bifurcation, posterior portion bifurcate anteriorly; posteriormargin of carapace with angular median prominence; intermediatecarinae of carapace not extending to anterior margin; posterolateralmargins not angled anteriorly.
Figure 8.
?
Squilla hesperia, new species, female holotype, TL 116.6, Baie de Tsimipaika:a, anterior portion of body; b, eye; c, outline of carpus of raptorial claw; d, outline oflateral processes of fifth, sixth, and seventh thoracic somites; e, last abdominal somite,telson, and uropod (setae omitted).Dactylus of raptorial claw with 6 teeth, outer margin of dactylussinuate; dorsal ridge of carpus with 3 tubercles; inferodistal angle ofmerus at most angled, not strongly spined.Mandibular palp and 4 epipods present.Last 3 thoracic somites with submedian and intermediate carinae,none armed; lateral process of fifth somite bilobed, anterior lobeproduced into a blunt, anteriorly directed spine, posterior lobe short,
no. 3G4i STOMATOPOD CRUSTACEA?MANNING 27
slender, triangular, directed laterally; lateral process of sixth somitebilobed, both lobes triangular, anterior lobe shorter and much slen-derer than posterior; lateral process of seventh somite bilobed, anteriorlobe obtuse, smaller than that of sixth somite, posterior lobe as longas but broader than posterior lobe of sixth somite; ventral keel ofeighth somite low, rounded.Abdomen with 8 carinae on first 5 somites, 6 on last, submedianssubparallel on each somite; second to fifth somites with anterior pairof tubercles lateral to intermediate carinae; abdominal carinae spinedas follows: submedian, 4-6; intermediate, 3-6; lateral, 1-6; marginal,1-5; sixth somite with sharp ventral spine on each side in front ofarticulation of uropod.Telson slightly longer than broad, with 6 sharp marginal teeth, eachwith short dorsal carina; prelateral lobe present; anterior portion ofmedian carina flanked by 2 short, oblique carinae at level of sub-median carinae of sixth somite; median carina interrupted anteriorly,terminating posteriorly in slender spine overhanging 2 tubercles;convergent line of pits lateral to median carina marked by mesialtubercles at middle of telson, mesial carina distally; remainder ofdorsal surface of telson smooth, convergent rows of pits neither inwell-marked grooves nor separated by carinae or low ridges; denticlesrounded, subequal, 3, 8, 1, each with proximal dorsal swelling; ventralsurface of telson with postanal keel.Proximal segment of uropodal exopod with 7-8 graded, movablespines, last short, not extending to midlength of distal segment; lobeon outer margin of inner spine of basal prolongation rounded, pro-jecting slightly, margin concave.Color.?Antennal scale outlined in dark pigment; carinae of bodydark; last 3 thoracic and first 5 abdominal somites with dark posteriorfine; telson with dark bar across surface, distal portion of mediancarina and marginal carinae dark; distal half of proximal segment andinner half of distal segment of uropodal exopod black; distal half ofuropodal endopod black.The color pattern is very similar to that found in S. massavensisKossmann.Size.?Female holotype, only specimen examined, TL 116.6. Othermeasurements: carapace length 23.3; cornea width 6.4; rostral platelength 4.0, width 4.0; telson length 21.1, width 20.2.Discussion.?Squilla hesperia closely resembles S. inassavensisKossmann, from the Red Sea but differs in having the rostral plateshorter and broader, in having the submedian carinae of the abdomendivergent on each somite rather than subparallel, and in having manyless dorsal tubercles on the telson. Squilla massavensis has two sub-median rows of tubercles converging distally on the median carinae of
28 PROCEEDINGS OF THE NATIONAL MUSEUM vol. imthe telson and has one or more tubercles lateral to these rows. Inaddition, adults of S. massavensis may have the dorsal surface of thetelson ornamented with curved rows of carinae between the curvedrows of pits; most of the dorsal surface of the telson in S. hesperiais smooth.In the holotype of S. hesperia the anterior lobe of the lateral processof the sixth thoracic somite is slenderer than that found in any of thespecimens of S. massavensis examined.Judging from the accounts of Kemp (1913) and Serene (1954),specimens identified as S. massavensis from outside of the Red Seado not belong to that species. I have seen small specimens of a Squillafrom the Persian Gulf in which the submedian carinae of the fourthabdominal somite are unarmed and which also differ from S. massaven-sis in having dark dorsal patches on the second and fifth abdominalsomites. It seems likely that specimens reported in the literature asS. massavensis from localities outside of the Red Sea belong to S.hesperia or to one or more undescribed species.The records of Miers (1880), Kemp (1913), and Ingle (1963), allbased on the same two specimens from Zanzibar, are included tenta-tively in the synonomy of S. hesperia; these two specimens must bereexamined.Etymology.-?The name is derived from the Latin, "hesperius,"meaning "western."Distribution.?-Known only from the type-locality.Squilla mauritiana Kemp, 1913Figure 9Squilla mauritiana Kemp, 1913, p. 68.Squilla juxtaoratoria Ward, 1942, p. 55.Previous records.?None.Material.? 1 9, 102.0; lie Europa, off Madagascar; P. Four-manoir; USNM.Description.?Eye of moderate size, cornea bilobed, set obliquelyon stalk; eyes not extending to end of first segment of antennularpeduncle; anterior margin of ophthalmic somite rounded, faintlyemarginate along midline; ocular scales obliquely truncate; cornealindex 397.Antennular peduncle shorter than carapace; dorsal processes ofantennular somite triangular, apices acute, directed anterolaterally.Antennal scale slender, curved, about three-fifths as long as cara-pace.Rostral plate slightly broader than long, appearing elongate, withupturned lateral margins; apex rounded; median tubercle present ondorsal surface.
STOMATOPOD CRUSTACEA?MANNING 29Carapace narrowed anteriorly, anterior width less than one-halfmedian length; median carina low, entire, not as sharp as remainderof carinae, with well-marked anterior bifurcation; distance fromdorsal pit to bifurcation less than distance from bifurcation to anteriormargin; portion of median carina posterior to cervical groove alsobifurcate anteriorly; posterior margin projecting along midline;4 lUl
Figure 9.
?
Squilla mauritiana Kemp, female, TL 102.0, He Europa: a, anterior portion ofbody; b, eye; c, carpus of raptorial claw; d, outline of lateral processes of fifth, sixth, andseventh thoracic somites; e, basal prolongation of uropod, ventral view.
anterolateral spines strong but not extending to base of rostral plate;intermediate carinae not extending to anterior margin; posterolateralportion of margin lacking anterior angle.Dactylus of raptorial claw with 6 teeth, outer margin sinuate;dorsal ridge of carpus with 2 tubercles; inferodistal angle of meruswith blunt spine.
30 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124Mandibular palp and 4 epipods present.Last 3 thoracic somites with unarmed submedian and intermediatecarinae; lateral process of fifth somite bilobed, anterior lobe a slender,anteriorly curved spine, posterior lobe slender, short, directed later-ally; lateral process of sixth somite bilobed, both portions triangular,acute, anterior much slenderer and shorter than posterior; lateralprocess of seventh somite bilobed, anterior lobe small, obtuse, posteriorlobe much larger, triangular; ventral keel of eighth somite rounded,inclined posteriorly.Abdomen with 8 carinae on first 5 somites, 6 on last, submediansslightly divergent on each somite; abdominal carinae spined asfollows: submedian, 4-6; intermediate, 2-6; lateral, 1-6; marginal,1-5; sixth somite with spine on each side in front of articulation ofuropod.Telson longer than broad, with 6 sharp marginal teeth, each withshort dorsal carina; prelateral lobe present; denticles rounded, 4-5,7-8, 1 ; ventral surface with postanal keel.Proximal segment of uropodal exopod with 9 graded, movablespines, last short, not extending to midlength of distal segment; lobeon outer margin of inner spine of basal prolongation prominent,rounded, margin concave.Color.?Largely faded in specimen from Madagascar except fordark pigment on uropods; proximal segment of exopod with distalhalf dark, distal segment with dark patch on proximal half of innerside; distal half of endopod dark.Size.?Only specimen examined, female, TL 102.0; other measure-ments of female: carapace length 23.0; cornea width 5.8; rostral platelength 3.5, width 3.8; telson length 21.4, width 18.9.Discussion.?In his account of the variation in Sguilla oratoria,Kemp (1913, p. 68) noted that two specimens from Mauritius differedfrom typical S. oratoria in having (a) a longer rostral plate, (b) theanterior portion of the lateral process of the sixth thoracic somiteshorter than the posterior, and (c) the submedian carinae of the fourthand the lateral carinae of the first and second abdominal somitesspined posteriorly. It was in this account that the name, "S. mauriti-ana," a manuscript name of Wood-Mason, was introduced. The speci-mens reported here differs from S. oratoria in these three features.In 1942 Ward described Sguilla juxtaoratoria from Mauritius. Al-though he gave some differences between his species and S. oratoria,his account was too brief to permit recognition of the species. Throughthe kindness of M. Claude Michel of the Mauritius Institute, I wasable to examine Ward's type and confirm that it is conspecific withthe specimen reported here. A more complete account of the type isin preparation in a review of the species of the uoratoria group."
N0
-
36" STOMATOPOD CRUSTACEA?MANNING 31Kemp (1913, p. 68) also noted that his specimens from Mauritiusresembled material from Hawaii, which shared the distinctive featuresmentioned above. The form from Hawaii is another distinct speciesthe description of which is in preparation.Sguilla mauritiana agrees with S. oratoria in having the mediancarina of the carapace uninterrupted; that character will distinguishboth species from S. fabricii Holthuis, S. inornata Tate, S. interruptaKemp, and S. perpensa Kemp. Sguilla woodmasoni, S. hesperia, andS. massavensis, other species of the "oratoria complex" that occur inthe western Indian Ocean, lack the anterior bifurcation on the mediancarina of the carapace; S. woodmasoni has a much broader carapaceand smoother telson than either S. hesperia or S. massavensis. Manning(1968) outlined the diagnostic features of several of the species in the
"oratoria complex."Distribution.?Indian Ocean, from Mauritius, and Madagascar.Squilla nepa Latreille, 1825Figure 10Squilla nepa.?Lenz, 1910, p. 571.?Kemp, 1913, p. 60, pi. 4 (fig. 49) [olderreferences].? Gravier, 1920, p. 377.?Holthuis, 1941, p. 245.?Barnard,1950, p. 847, figs, lb, 2a.?Fourmanoir, 1953, p. 153.?Kurian, 1954, p. 85?Serene, 1954, pp. 6, 8.?Stephenson and McNeill, 1955, p. 243.?Crosnier,1965, p. 61 [listed].Previous records.?Tamatave (Lenz, 1910); Diego Suarez(Gravier, 1920); several localities (Fourmanoir, 1953); no specificlocality (Crosnier, 1965).Material.?2 9, 122.6-127.0; Baie Narendry, northwesterncoast; trawl; 6-7 m; soft mud; A. Crosnier; February 1958; MNHNP.2 9, 85.8-128.6; same; USNM. 5 d", 89.6-132.2; 4 9, 100.7-135.8;Ambaro Bay, Nosy Be; trawl; 2-5 m; M. Pichon; 8 September 1964;USNM. 12 d", 89.2-126.7; 24 9, 70.0-146.0 (in 5 lots) ; from localitiesaround Nosy Be; J. Rudloe, col.; IIOE; January-February 1964;USNM.Diagnosis.?Eye very small, cornea set transversely on stalk;corneal indices 514-690; rostral plate quadrangular or subtriangular;anterolateral spines of carapace strong; median carina of carapacewith anterior bifurcation open for half or more of its length; dactylusof raptorial claw with 6 teeth, outer margin of dactylus sinuate;dorsal ridge of carpus with 2-3 tubercles; inferodistal angle of merusof claw armed; mandibular palp and 4 epipods present; lateral pro-cesses of fifth to seventh thoracic somites bilobed; abdominal carinaespined as follows: submedian, (3) 4-6; intermediate, (2-3) 4-6;lateral, 1-6; marginal, 1-5; denticles, 1-3, 7-9, 1, rounded; ventralsurface of telson with postanal keel.
32 PROCEEDINGS OF THE NATIONAL MUSEUMColor.?Entire body dusky, appearing mottled; carinas green inpreservative; some specimens with transverse patch of dark pigmenton second abdominal somite; distal portions of uropod dark, exceptfor distal segment of exopod, which is light with dark edges.Discussion.?The present specimens agree well with accounts ofthe species in the literature. In shape of rostral plate and spinationof the abdomen they show an unusual amount of variation. The
Figure 10.
?
Squilla nepa Lacreille, female, TL85. 8, Baie Narendry : anterior portion of body.
rostral plate may be triangular or subquadrate, with the apex trans-verse or rounded. In one specimen the plate is very short and broadlyrounded. The submedian carinae of the third abdominal somiteare armed in about half the specimens; the spines may be present onlarge or small specimens. The intermediate carinae of the secondand third somites may be armed.Distribution.?Western portion of Indo-West Pacific region,from the western Indian Ocean and South Africa to Hong Kong.
Acanthosquilla Manning, 1963Four species of Acanthosquilla have been recorded from the Indo-West Pacific region and one new species is described herein. Thefollowing key will distinguish the five species, of which all but one,A. tigrina (Nobili), occurs in the western Indian Ocean.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 33Key to Indo-West Pacific Species of Acanthosquilla
1. Telson with 4 pairs of fixed marginal teeth 2Telson with 2 pairs of fixed marginal teeth 42. Ventral margin of sixth abdominal somite with posterior spines.A. tigrina (Nobili, 1903)Ventral margin of sixth abdominal somite unarmed 33. Rostral plate trispinous; dactylus of claw with 10-11 teeth; submediandenticles arranged in semicircle A. vicina (Nobili, 1904)Rostral plate angled anterolateral^, with median spine; dactylus of claw with7 teeth; submedian denticles in transverse row . . A. humesi, new species4. Lobes on outer margin of dactylus subequal; submedian denticles in trans-verse row; first and third intermediate denticles larger than second andfourth A. acanthocarpus (Miers, 1880)Distal lobe on outer margin of dactylus much larger than proximal; submediandenticles in oblique row; second and fourth intermediate denticles largerthan first and third A. multifasciata (Wood-Mason, 1895)
Acanthosquilla humesi, new speciesFigure 11Acanthosquilla Humes, 1965, p. 184.Holotype.? 1 9, 62.5; Ambatozavary, Nosy Be, Madagascar;E. Cutler; 16 July 1964; USNM 124095.Paratypes.? 1 d\ 63.2; 1 9, 64.5; Antsakoabe, northwestern shoreof Nosy Be, Madagascar; dug from 30 cm intertidal sand; A. G.Humes; host no. 889; 12 July 1964; USNM 124096.Description.?Eye small, not extending to end of antennularpeduncle; cornea subglobular, slightly larger than and set obliquelyon stalk; ocular scales small, erect, bases fused, apices divergent.Antennular peduncle short, less than half as long as carapace;dorsal processes of antennular somite visible on either side of rostralplate as a slender, anteriorly directed spine.Antennal peduncle with 1 mesial and 1 ventral papilla; antennalscale short, less than half as long as carapace.Rostral plate trapezoidal, narrowed anteriorly, angled antero-laterally, with slender median spine; anterior portion of median spineresting in apex of V formed by ocular scales.Carapace narrowed anteriorly, rounded anterolaterally and poste-riorly, lacking spines, carinae, or cervical groove.Raptorial claw stout, dactylus with 7 teeth, penultimate shorterthan antepenultimate; base of dactylus with proximal angled projec-tion and distal obtuse projection separated by a shallow concavity;propodus with 4 proximal movable spines and normal pectination;dorsal ridge of carpus terminating in slender spine; merus much longerthan ischium.273-102?67 3
34 PROCEEDINGS OF THE NATIONAL MUSEUMPropodi of third and fourth thoracic appendages broader than long,with ventral ribbing; propodus of fifth appendages as broad as long!with inferior brush of setae.Mandibular palp and 5 epipods present.Exposed thoracic somites smooth, lateral margins rounded or sub-truncate; basal segment of each walking leg unarmed; endopods of
,*? ,' .j 's$? a
w <&x m*.Figure \\.?Acanthosquilla humesi, new species, female holotype, TL 62.5, Nosy Be: aanterior portion of body; b, raptorial claw; c, last two abdominal somites, te'lson, and uro-pod; d, telson, ventral view; e, uropod, ventral view (setae omitted).walking legs 2-segmented, subcircular on first 2 legs, ovate on last;eighth somite without noticeable ventral keel.Abdomen depressed, somites broad, smooth, unarmed except forslender posterolateral spines on sixth; sixth somite with prominent
no. 3641 STOMATOPOD CRUSTACEA?MANNING 35
ventral projections overhanging articulation of uropod, each armedwith accessory spinule; ventral surface of sixth abdominal somiteunarmed.Telson broader than long, with normal dorsal fan of 5 teeth; mar-ginal armature consisting of, on either side of the midline, 1 movablesubmedian tooth and 4 fixed teeth, inner overhanging movable sub-medians; 6-7 submedian denticles present, with 1 denticle presentbetween each of the other marginal teeth.Uropod with 7 slender movable spines present on outer margin ofproximal segment of exopod, last extending to midlength of distalsegment; 9-13 stiff setae present on inner distal lobe of proximalsegment of exopod; endopod triangular, with prominent fold oninner, proximal margin; spines of basal prolongation trefoil in crosssection, inner longer than outer.Color.?Kostral plate, eyestalks, anterior appendages, and dorsalsurface of claws with scattered dark spots; carapace with 3 transverseblack bands plus dark crescent at each posterolateral angle; last 3thoracic and all 6 abdominal somites with broad, transverse blackband; telson with 4 oval dark patches, interrupted at midline andbetween second and third dorsal teeth on each side; uropodal exopodwith black spot at articulation of distal segment, inner angle of distalsegment dark; endopod black.Size.?Only male known, TL 63.2; females, TL 62.5-64.5. Othermeasurements of female paratype: carapace length 10.6; cornea width1.6; rostral plate length 3.1, width 3.4; fifth abdominal somite width11.1; telson length 5.0, width 8.8.Discussion.? A. canthosquilla humesi can be distinguished fromboth A. acanthocarpus (Miers) and A. multifasciata (Wood-Mason)by the presence of four fixed marginal teeth on the telson; the lattertwo species both have but two fixed spines. Acanthosquilla humesiagrees with A. tigrina (Nobili) and A. vicina (Nobili) in having fourpairs of fixed marginal teeth on the telson but may be readily dis-tinguished from both of those species. It lacks the posterior ventralspines on the sixth abdominal somite that are characteristic of A.tigrina, and it differs from A. vicina in having a single median spineon the rostral plate, fewer spines on the claw (seven instead of 10-11),and in having the submedian denticles of the telson arranged in atransverse row rather than in a semicircle.These specimens were the hosts of a cyclopoid copepod, Hemicyclopsacanthosquillae Humes, collected and described by Arthur G. Humes(1965). This, I believe, is the first recorded occurrence of a copepod-stomatopod association.Humes (1965, p. 184) noted that the stomatopods appeared in thewater seeping into a hole 30 cm deep that had been dug in clear sand.
36 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124Etymology.?The species is named for the collector, Arthur G.Humes.Distribution.?Known only from Nosy Be, Madagascar.Lysiosquilla Dana, 1852Only two species of Lysiosquilla have been recognized from theIndo-West Pacific region in recent years, L. capensis Hansen, 1895,and L. maculata (Fabricius, 1793). Two varietal forms of the latterspecies, L. m. var. sulcirostris Kemp, 1913 and L. m. var. tredecim-dentata Holthuis, 1941, also have been recognized. All four of theseforms are distinct species that may be distinguished by means of thekey below.A redescription of L. capensis Hansen is in preparation.Two species of Lysiosquilla, maculata and tredecimdentata, occur offMadagascar.Key to Indo-West Pacific Species of Lysiosquilla1. Rostral plate triangular, greatest width at base; median carina of plateflanked by longitudinal grooves; dactylus of claw with 8 teeth.L. sulcirostris Kemp, 1913Rostral plate cordiform, greatest width in advance of base; median carina ofplate, when present, not flanked by longitudinal grooves; dactylus of clawwith more than 8 teeth 22. Antennal scale oval, less than twice as long as broad; anterior margin ofantennal protopod lacking projection; ventral keel of eighth thoracicsomite rounded L. maculata (Fabricius, 1793)Antennal scale slender, elongate, more than twice as long as broad; anteriormargin of antennal protopod with projection; ventral keel of eighth tho-racic somite acute, sharp, posteriorly 33. Rostral plate lacking median carina; ventral surface of uropodal protopodwith slender spine at articulation of endopod . . L. capensis Hansen, 1895Rostral plate with median carina; ventral surface of uropodal protopod lack-ing spine at articulation of endopod . . L. tredecimdentata Holthuis, 1941Lysiosquilla maculata (Fabricius, 1793)Figure 12Lysiosquilla (Miers) maculata.?Lenz and Richters, 1881, p. 428.Lysiosquilla maculata.?Kemp, 1913, p. Ill, pi. 8 (fig. 89-91) [older references].
?
Monod, 1925, pi. xxi (fig. d).?Holthuis, 1941, p. 269, fig. 5.?Barnard, 1950,p. 855, figs. 3d, 4c-d?Serene, 1954, pp. 6, 8, 11, 13, 64, 66, fig. 12, pi. 5(figs. 1-2), pi. 6 (figs. 1-2).?Kurian, 1954, p. 86.?Stephenson and McNeil,1955, p. 246.?Manning, 1962, p. 2; 1967, p. 103.Lysiosquilla maculata maculata.?Ingle, 1963, p. 23, figs. 23, 45, 61, 72 [part; notspecimen from Gulf of Aden].Previous records.?Lenz and Richters, 1881; Monod, 1925.Material.? 1 9, 146.0; Nosy B6, Madagascar; intertidal zone; A.Crosnier; May 1960; USNM.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 37Description.?Eye large, cornea bilobed, set almost transverselyon stalk; eyes not extending beyond end of second segment of anten-nular peduncle; ocular scales triangular, apices acute, curved anteri-orly; corneal index 362.Antennular peduncle short, about half as long as carapace; dorsalprocesses of antennular somite visible lateral to rostral plate asanteriorly directed spines.Antennal scale broad, ovate, more than twice as long as broad;antennal peduncle extending to or slightly beyond eye; basal portionof antenna with anterior margin straight, not produced into ananterior triangular projection or spine, and with 3 papillae, 1 mesial,2 ventral.Rostral plate cordiform, broader than long, apex deflexed; mediancarina present on anterior half.
Figure 12.
?
Lysiosquilla maculata (Fabricius), female, TL 146.0, Nosy Be: a, antennalscale (setae omitted); b, color pattern of ischium and merus of raptorial claw; c, outlineof ventral keel of eighth thoracic somite.
Dactylus of raptorial claw with 9-11 teeth (10 in present specimen),outer margin of dactylus sinuate; propodus with few hairs proximally;dorsal ridge of carpus terminating in single spine, apex not deflexed.Mandibular palp and 5 epipods present.Ventral keel of eighth thoracic somite low, rounded, unarmed.Abdomen smooth, unarmed; sixth somite with irregular submedianarea separated from smooth lateral boss by a longitudinal sulcus; sixthsomite with slender, triangular projection in front of articulation ofeach uropod.Telson much broader than long, pitted dorsally, with low, raisedtriangular median boss and 2 lower, oval, submedian bosses on dorsalsurface; posterior margin with 4 pairs of projections, outer 2 bestdeveloped.
38 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124Basal segment of uropod with dorsal spine; proximal segment ofexopod with 8 slender, movable spines, last not extending to midlengthof distal segment; distal segment of exopod longer than proximal;endopod broad, triangular, twice as long as broad; basal prolongationconsisting of 2 spines, trefoil in cross section, inner longer; ventralsurface of uropodal protopod unarmed at articulation of endopod.Color.?Body with dark bands, pattern variable (see illustrationin Kemp, 1913); carapace with 3 dark bands, body with dark band atarticulation of each somite; antennal scale with 1-2 dark patches, notoutlined with dark pigment; claw conspicuously banded with darkpigment; basal portion of first walking leg light, of last 2 dark; sixthabdominal somite light dorsally; telson with median and 2 submediandark parches, connected anteriorly; basal segment of uropod darkproximally; distal half of distal segment clear; most of distal portionof endopod dark.Size.?Only specimen examined, female, TL 146.0. Other measure-ments: carapace length 26.4; cornea width 7.3; rostral plate length 5.3,width 5.9; antennal scale length 16.5, width 7.1; fifth abdominalsomite width 30.5; telson length 20.2, width 28.6.Discussion.?The unarmed antennal and uropodal protopods,broad rostral plate, broad antennal scale, and characteristically barredclaw will distinguish this species from others in the genus.Distribution.?Widely distributed in the Indo-West Pacific, fromnumerous localities between the western Indian Ocean and Hawaii.Lysiosquilla tredecimdentata Holtliuis, 1941Figure 13?Lysiosquilla maculata.?Chopra, 1939, p. 16.Lysiosquilla maculata var. tredecimdentata Holthuis, 1941, p. 273, fig. 6.Lysiosquilla maculata tredecimdentata.?Manning, 1963, p. 317 [listed].?Lysiosquilla maculata maculata.?Ingle, 1963, p. 23 [part; reference to specimenfrom Gulf of Aden only].Previous records.?None.Material.? 1 cf, 161.7; Estuaire du Mangoky, western coast ofMadagascar; trawl; 9 m; grey mud, sand; A. Crosnier; June 1959;USNM.Description.-?Eye large, cornea bilobed, set almost transverselyon stalk; eye not extending beyond end of second segment of anten-nular peduncle; ocular scales subtriangular, flattened dorsally, apicesacute, directed anteriorly; corneal index 354.Antennular peduncle short but more than half as long as carapace;dorsal processes of antennular somite visible lateral to rostral plateas anteriorly directed spines.
STOMATOPOD CRUSTACEA?MANNING 39Antennal scale slender, curved, more than 3 times as long as broad;antennal peduncle not extending beyond eye; basal portion of antennawith anterior triangular projection, directed anterolaterally, and with3 papillae, 1 mesial, 2 ventral.
Figure 13.
?
Lysiosquilla tredecimdentata Holthuis, male, TL 161.7, Mangoky estuary: a,anterior portion of body; b, color pattern of ischium and merus of raptorial claw; c,outline of ventral keel of eighth thoracic somite; d, last abdominal somite, telson, anduropod. (Setae omitted).
40 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124Rostral plate cordiform, broader than long, apex deflexed; mediancarina present on anterior half.Raptorial claw with 10 teeth (12-13 on other specimens), outermargin of dactylus sinuate; propodus with some proximal hairs, as infemale L. maculata; dorsal ridge of carpus terminating in a singlespine, deflexed mesioventrally.Mandibular palp and 5 epipods present.Ventral keel of eighth thoracic somite produced into a posteriorlydirected spine.Abdomen smooth, unarmed; sixth somite with irregular submedianarea separated from longitudinal lateral boss by a longitudinal sulcus;sixth somite with blunt triangular projection in front of articulationof each uropod.Telson much broader than long, pitted dorsally, with low, raisedtriangular median boss and 2 lower oval submedian bosses on dorsalsurface; posterior margin with 4 pairs of projections, outer 2 bestdeveloped.Basal segment of uropod with dorsal spine; proximal segment ofexopod with 9 slender, movable spines on outer margin, last notextending to midlength of distal segment; distal segment of exopodlonger than proximal; endopod broad, triangular, over twice as longas wide; basal prolongation consisting of 2 spines, trefoil in crosssection, inner longer; ventral surface of uropodal protopod unarmedat articulation of endopod.Color.?Body with dark bands, as in L. maculata; carapace with3 dark bands and body with broad, dark band at articulation of eachsomite; claw lacking prominent dark bars, distal half of merus andproximal half of propodus with dark flush; basal fourth of all walkinglegs dark; sixth abdominal somite dark; telson with broad, transversedark patch, most of distal margin lighter; basal segment of uropoddark; distal half of proximal segment and proximal half of distalsegment of uropodal exopod dark, distal half of distal segment light;most of distal portion of uropodal endopod dark.Size.-?-Single male examined, TL 161.7. Other measurements:carapace length 28.3; cornea width 8.0; rostral plate length 6.4,width 7.4; antenna! scale length 17.2, width 5.4; fifth abdominalsomite width 35.2; telson length 21.8, width 32.0.Discussion.-?The slender antennal scale, outlined in dark pigment,the anterior projection on the basal portion of the antenna, the lackof dark bands on the claw, the deflexed spine on the carpus of the claw,the triangular keel on the eighth thoracic somite, and the unarmedventral surface of the uropodal protopod will distinguish this speciesfrom all others in the genus.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 41Lysiosquilla tredecimdentata has not been recorded previously fromoutside the Gulf of Aden. I have compared this specimen, a male,with three females taken at Aden Harbor during the InternationalIndian Ocean Expedition. It differs in having a larger eye, which isbroader than the rostral plate, and in having fewer teeth, 10 insteadof 12, on the dactylus of the claw. I have assumed that these aresexual differences, for in all other respects the four specimens arealmost identical, even to overall color pattern.Chopra (1939) noted that his specimen of L. maculata from theGulf of Aden had 12 teeth on the dactylus of the claw, an unusuallyhigh number for L. maculata. All the specimens of L. tredecimdentatafrom the Gidf of Aden examined by me had 12 teeth on the claw.It seems likely that Chopra was dealing with L. tredecimdentata notL. maculata. Ingle (1963) also listed the occurrence of the samespecimen previously recorded by Chopra.The single male specimen from Madagascar shows some trace of thehairs on the propodus of the raptorial claw characteristic of adultfemales of L. maculata. There is no apparent reduction in size of theclaw or in size and number of teeth.Distribution.?Western Indian Ocean, from Aden (?Chopra,1939; Holthuis, 1941; ?Ingle, 1963) and from Madagascar.Odontodactylus Bigelow, 1893Three species of this genus are known to occur in the Indian Ocean,and two of these are recorded below from Madagascar. Odontodactylusbrevirostris (Miers) , which is not represented in the present collection,has been reported from the Seychelle Islands and from ProvidenceIsland (Miers, 1884; Borradaile, 1907; Manning, 1967a) and could beexpected off Madagascar.The three specimens reported below were included by me in areview of the genus Odontodactylus, which contained complete ac-counts for each of the five known species.Odontodactylus japonicus (de Haan, 1844)Odontodactylus japonicus.?Kemp, 1913, p. 139.?Manning, 1967a, p. 7, fig. 2[other references].Illustration.?Manning, 1967a, fig. 2.Previous records.?Ambovombe (Manning, 1967a).Material.? 1 9, 117.0; Ambovombe, southern coast of Madagas-car; 60 m; shelly sand; A. Crosnier; 8 October 1958; USNM.Remarks.?This specimen was recorded by Manning (1967a).Distribution.?Indo-West Pacific, from two localities in theIndian Ocean, Seychelles (Borradaile, 1907) and Madagascar (Mann-
42 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124ing, 1967a), and from locations around China and Japan. In moderatedepths, to 200 m.Odontodactylus scyllarus (Linnaeus, 1758)Gonodactylus scyllarus.?Miers, 1880, p. 115.? Richters, 1880, p. 167.Odontodactylus scyllarus.?Kemp, 1913, p. 135.?Manning, 1967a, p. 10, fig. 3[other references].Illustration.?Manning, 1967a, fig. 3.Previous records.?Miers, 1880; Richters, 1880; Manning, 1967a.Material.? 1 9, 131.0; Nosy Be, Madagascar; P. Fourmanoir;MNHNP. 1 broken tf , CL 36.0; same; A. Crosnier; USNM.Remarks.?These specimens were reported by Manning (1967a).Distribution.?Indo-West Pacific, from the western Indian Oceanto Japan. Shallow water.Pseudosquilla Dana, 1852Pseudosquilla ciliata (Fabricius, 1787)Pseudosquilla ciliata.?Lenz, 1910, p. 571.?Kemp, 1913, p. 96 [older references].
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Holthuis, 1941, p. 261 [older references].?Barnard, 1950, p. 852, fig. 3a?Serene, 1951, p. 11, figs 1-6, 8-i; 1954, pp. 6, 10.?Stephenson and McNeill,1955, p. 245.?Manning, 1962, p. 2.?Ingle, 1963, p. 21, figs. 18, 39, 52, 68.Illustration.?Serene, 1951, figs. 1-6, 8-1.Previous records.?Sainte Marie (Lenz, 1910); Fort Dauphin(Gravier, 1935).Material.? 1 9, 36.8; Nosy B6, Madagascar; intertidal zone; A.Crosnier; September 1958; USNM. 1 cf, 37.7; Nosy lava (?); A.Crosnier; MNHNP. 1 cf , 37.8; 1 9, 24.2; Banc de Pracel, westerncoast of Madagascar; 55 m; muddy sand; June 1959; USNM. 1 9,80.3; Anjouan Id., Comoro Ids.; A. Crosnier; USNM. 1 9, 63.8Anjouan Id., Comoro Ids.; from stomach of Caranx; A. CrosnierMNHNP. 1 postlarva, 17.9; Mayotte Id., Comoro Ids.; 47 m; sandA. Crosnier; August 1959; MNHNP. 1 9, 20.0; same; 51 m; coarsesand; A. Crosnier; 1 October 1959; MNHNP.Remarks.?All specimens correspond to the "forme claire" dis-cussed by Serene (1951).Distribution.?Widely distributed in the Atlantic and Indo-WestPacific Oceans. Gonodaclylus Berthold, 1827The Indo-West Pacific species of Gonodactylus fall into three dis-tinct groups or sections, one centered around 67. chiragra (Fabricius),one around G.jalcatus (Forskal), and one around 67. demanii Henderson.The "chiragra section" includes large species with broad ocularscales and with three large carinae on the median dorsal surface of the
no. 3641 STOMATOPOD CRUSTACEA?MANNING 43
telson. Species in this section include the three species reported below,G. chiragra (Fabricius), G. platysoma Wood-Mason, and G. smithiiPocock. It is suggested below that G. acutirostris de Man is based onan aberrant specimen, and that de Man's species is conspecific withG. smithii.The "falcatus section" includes only two species, G. falcatus (For-skal) and G. graphurus Miers; these species have small ocular scalesand five carinae on the median area of the telson. Manning (1966)and others have pointed out that the latter species does not occurwest of Indo-Malaya and that earlier records to it in the literaturefrom the western Indian Ocean are referable to G. falcatus.The "demanii section" includes 10 small species, mature at TL50 mm or less, seven of which were reported by Manning (1967b).Three additional species are reported below from Madagascar, in-cluding one species revived from the synonoymy of G. chiragra andtwo described as new.Gonodactylus chiragra (Fabricius, 1781)Gonodactylus chiragra.?Lenz and Richters, 1881, p. 428.?Kemp, 1913, p. 155,fig. 2 on p. 161, pi. 9 (fig. 107).? Gravier, 1920, p. 377; 1935, p. 358?Holthuis, 1941, p. 277, fig. 7 [older references].?Poisson, 1949, p. 23 ?Barnard, 1950, p. 861.?Fourmanoir, 1952, p. 171; 1953, p. 157.?Serene,1954, pp. 6, 10, 11, 27, 42, 74, 84, figs. 9-10, 13-1, 13-2, 15, pi. 7.?Stephensonand McNeill, 1955, p. 250.?Manning, 1965, p. 2; 1966, p. 113; 1967, p. 102.Not Gonodactylus chiragra chiragra.?Ingle, 1963, p. 27, figs. 27, 47, 63. [=G.smithii Pocock?]Illustration.?De Man, 1898, pi. 38 (fig. 77) ; Kemp, 1913, fig. 1on p. 161.Previous records.?Fort Dauphin (Gravier, 1935) ; He Europa(Fourmanoir, 1952); no specific locality (Lenz and Richters, 1881;Gravier, 1920; Poisson, 1949; Fourmanoir, 1953).Material.? 1 broken 9, CL 13.9; Mayotte Island, Comoro Islands;reef flat; L. S. Kornicker, Sta. LK-39; Anton Bruun Cruise 9, IIOE;24 November 1964; USNM. 1 9, 48.3; Nosy Be\ Madagascar; A.Crosnier; 10 December 1958; MNHNP. 1 d\ 83.5; Nosy B6; underdead coral debris; Mme. Chavane; 18 September 1958; MNHNP.1 d" , 70.9; 2 9, 48.4-68.9; Nosy B6; in sandstone; 20 May 1958;MNHNP. 4 d\ 31.8-55.7; 5 9, 22.8-72.1; Nosy B6; sandstone;A. Crosnier; November 1961; USNM. 1 cf , 70.5; Ambatoloaka, NosyBe; Mme. Chavane; 25 September 1964; USNM. 8 cf , 22.4-88.4;3 9, 41.9-67.3; Pt. Fievre, Nosy Be; 13?24'21" S, 48?18'33" E;J. Rudloe, field no. Jr-13; IIOE; 26 December 1963; USNM. 2 d\50.3-51.2; Ambatomboka (?) Crater Point, Nosy B6; 13?24'17" S,48?13'31" E; taken from base of volcanic rocks; J. Rudloe, Field no.JR-23; IIOE; 3 January 1964; USNM. 1 d\ 44.1; 1 9, 51.6; 50 yds.
44 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121
right of Centre d'Oceanographie pier, Pt. Fievre, Nosy Be; 13?24.4'S, 4S?17.8 / E; intertidal mud flat, mud, rock; J. Rudloe, field no.24A; IIOE; 4 January 1964; USNM. 1 d\ 105.5; mud flats of Amba-tozavavy, Nosy Be; 13?22'10" S, 48?19'52" E; J. Rudloe, field no.JR-52; IIOE; 29 January 1964; USNM. 1 d" , 42.7; Tani Keli (TanyKely), Nosy Be; 13?26'36" S, 48?14'48" E; J. Rudloe, field no.JR-62; IIOE; USNM. 1 d" , 21.3; 3 9, 68.5-73.2; Nosy Be; J. Rudloe,field no. JR-63-1; IIOE; 17 December 1963; USNM. 1 <?, 51.3; 2 9,67.4-69.2; Nosy Be; J. Rudloe, field no. JR-63-2; IIOE; USNM.Distribution.?Widely distributed in the Indo-West Pacific region,from the western Indian Ocean through Oceania.Gonodactylus platysoma Wood-Mason, 1895Gonodactylus chiragra var. acutus Lanchester, 1903, p. 447, pi. 23 (figs. 2, 3, 3a).
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Lenz, 1910, p. 571.Gonodactylus chiragra var. tumidus Lanchester, 1903, p. 447, pi. 23 (figs. 1, la).Lenz, 1910, p. 571.Gonodactylus chiragra var. platysoma.?Kemp. 1913, p. 162, fig. 1 on p. 161.
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Bigelow, 1931, p. 117, pi. 1 (fig. 2), pi. 2 (fig. 2) .?Holthuis, 1941, p. 281.Gonodactylus platysoma.?Serene, 1954, pp. 10, 74, fig. 13-4.?Manning, 1962,p. 3; 1966, p. 110; 1967, p. 103.Gonodactylus chiragra platysoma.?Ingle, 1963, p. 25 [key].Illustration.?Bigelow, 1931, pi. 1 (fig. 2), pi. 2 (fig. 2).Previous records.?Tamatave, Sainte Marie, Angontil (Lenz,1910).Material.? 1 d\ 72.8; Mayotte Id., Comoro Islands; intertidalzone; A. Crosnier; September 1959; MNHNP. 1 cf, 59.7; AnjouanId., Comoro Islands; intertidal zone; A. Crosnier; November 1961;USNM.Remarks.?The specimen from Anjouan Island has two commensalgastropods attached to the pleopods. Holthuis (1951) has commentedon the occurrence of a gastropod, Caledoniella montrouzieri Souverbie,on specimens of Gonodactylus chiragra. As far as I can determine,no commensals have been reported for 67. platysoma.Distribution.?Indo-West Pacific, from scattered localities be-tween the western Indian Ocean and Oceania.Gonodactylus smithii Pocock, 1893Gonodactylus smithii Pocock, 1893, p. 475, pi. 20b (fig. 1).?Serene, 1954, pp. 6>7, 10, 74, 76, fig. 13-5, pi. 8.?Manning, 1966, p. 112.Gonodactylus chiragra var. acutirostris de Man, 1898, p. 695, pi. 38 (figs. 77b, c).Lanchester, 1903, p. 454.? Gravier, 1937, p. 204, fig. 20.Gonodactylus chiragra var. smithii.?Borradaile, 1898, p. 33, 35.?Lanchester,1903, p. 447, pi. 23 (figs. 4-5).?Lenz, 1905, p. 387.?Tattersall, 1906,p. 167.?Borradaile, 1907, p. 211 [key], p. 212.?Lenz, 1910, p. 571.? Rath-bun, 1914, p. 664.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 45Gonodaclylus chiragra var. c {acutirostris).?Borradaile, 1899, pp. 400, 401, 402.Gonodactylus chiragra var. d (smithi).?Borradaile, 1899, p. 402.Gonodaclylus chiragra.?Lanchester, 1901, p. 555.? Nobili, 1906, p. 157 [part].Tattersall, 1921, p. 359 [smithii in discussion].Gonodactylus acutirostris.?Borradaile, 1907, p. 210 [key].?Kemp, 1913, p. 163.
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Serene, 1947, p. 382, fig. 1, pi. 1.?Dawydoff, 1952, p. 145.?Serene, 1953,pp. 506, 507.Gonodactylus chiragra var. intermedia de Man, 1929, p. 25, pi. 3 (figs. 9-9b);1929a, p. 3.f Gonodactylus chiragra chiragra.?Ingle, 1963, p. 27, figs. 27, 47, 63 [part?].Illustration.?Serene, 1947, fig. 1, pi. 1.Previous records.?None.Material.? 1 cf, 38.9; Mayotte Id., Comoro Islands; reef flat;L. S. Kornicker, Sta. LK-39; Anton Bruun Cruise 9, IIOE; 24 No-vember 1964; USNM. 1 9, 29.4; Anjouan Id., Comoro Islands;intertidal zone; A. Crosnier; November 1961; USNM. 1 9, 41.0; NosyB6, Madagascar; in sandstone; 20 May 1958; MNHNP. 1 9, 30.2;Banc de FEtoile, Madagascar; dredge; 20-80 m; A. Crosnier; May1960; MNHNP. 1 c/\34.7; 1 9, 41.7; Tulear, Madagascar; A. Crosnier;October 1958; USNM. 1 d\ 50.0; 2 9, 44.4-48.2; Tulear; from off-shore reef; K. J. Boss; Anion Bruun Cruise 7, IIOE; 9 August 1964;USNM.Discussion.?I believe Lanchester (1903) was correct in suggestingthat G. acutirostris de Man was based on a specimen with a damagedtelson, with the deformity affecting the posterior end of the mediancarina and the anterior ends of the carinae of the submedian teeth.In other respects, including the sharpness of carinae on the telson andthe acute anterolateral angles of the rostral plate, it agrees with G.smithii Pocock. There is little doubt in my mind that G. chiragra var.intermedia de Man, from Pulau Berhala, is conspecific with G. smithii;it agrees with the latter species in all respects.Odhner (1923) pointed out that Kemp (1913), in his account ofG. acutirostris, misunderstood de Man's original description in statingthat G. acutirostris lacked the anterior tubercles on the telson; Odhnerpointed out that these tubercles were present, but that the type lackedthe flukes of the anchor. In my account of G. smithii (1966) fromAustralia, I made the same error in pointing out differences betweenmy specimens and G. acutirostris. De Man (1929) noted that in thetype of G. acutirostris the anterior tubercles were present.Specimens of G. smithii from Australia differ from those reportedhere in having a broader endopod on the uropod, on which the innermargin is convex. In the specimens from Madagascar the inner marginof the uropodal endopod is sinuous, convex proximally, concavedistally.
46 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124One of the male specimens from Tulear has two gastropod molluscsattached to the body, a large one on the pleopods and a smaller oneon the ventral surface of the eighth thoracic segment. As noted underG. platysoma, commensal gastropods have been reported only for67. chiragra.Distribution.?Indo-West Pacific, from the western Indian Oceanto Viet Nam and Australia. Shallow water to 50 fms.Gonodactylus falcatus (Forskal, 1775)Gonodactylus glabrous.?Kemp, 1913, pp. 167, 197, fig. 2 on p. 170, pi. 9 (fig. 113)[older references].?Barnard, 1950, p. 863, fig. 3f.Gonodactylus falcatus.?Holthuis, 1941, p. 284, fig. 9a.?Serene, 1954, pp. 6, 7, 10,11, 42, 77, figs. 8, 13-6, pi. 9.?Stephenson and McNeill, 1955, p. 249.?Manning, 1962, p. 4.?Ingle, 1963, p. 29, figs. 28, 57.?Manning, 1965, p. 260;1966, p. 109; 1967, p. 102.Illustration.?Brooks, 1886, pi. 14 (fig. 5).Previous records.?None.Material.? 1 9, 21.1; Mayotte Id., Comoro Islands, reef flat;L. S. Kornicker, Sta LK-39; Anton Bruun Cruise 9, IIOE; 24 Novem-ber 1964; USNM. 1 d\ 35.9; 1 9, 22.8; He Europa, off Madagascar;A. Crosnier; MNHNP. 1 6% 63.3; lies Mitsio, northwestern coast ofMadagascar; trawl; 26 m; sand; A. Crosnier; June 1959; MNHNP.2 d1 , 33.8-54.5; 11 9, 37.7-71.6; Sud Mitsio, northwestern coast ofMadagascar; trawl; 26 m; sand; A. Crosnier; June 1959; USNM.1 cf, 20.3; Banc de Pracel, western coast of Madagascar; 50 m;muddy sand; A. Crosnier; June 1959; MNHNP.Distribution.?Indo-West Pacific region, from Ked Sea and SouthAfrica to Japan. Gonodactylus bicarinatus, new speciesFigure 14Holotype.? 1 9, 26.7; Nosy B6, Madagascar; intertidal zone;A. Crosnier; October 1959; USNM 124097.Description.?Anterior margins of rostral plate straight, antero-lateral angles rounded.Ocular scales small, erect.Mandibular palp and 5 epipods present.Lateral processes of sixth and seventh thoracic somites subtruncate,rounded laterally, subequal.Carinae of sixth abdominal somite not markedly inflated, each withapical spine.Telson broader than long, with 3 pans of slender, sharp marginalteeth, laterals well developed; dorsal surface of telson with numerousspinules and tubercles, smaller and more abundant on anterior carinaethan on carinae of marginal teeth; median carina oval, very inflated,
STOMATOPOD CRUSTACEA?MANNING 47
with 1-2 irregular marginal rows of tubercles, apical spine largest,median surface smooth; anterior submedian carinae with 1-2 dorsalspinules, 1 distal spine, and distolateral patch of spinules; knob with3 spinules; anterior surface of telson with rounded lobe, each tuber-culate, at level of intermediate carinae of sixth somite; carinae ofsubmedian and intermediate marginal teeth each ornamented withnumerous large spines; numerous small submedian and 2 sharp inter-mediate denticles present; outer intermediate denticle recessed ante-
Figure 14.
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Gonodactylus bicarinatus, new species, female holotype, TL 26.7, Nosy Be: a,outline of rostral plate; b, outline of lateral processes of sixth, seventh, and eighth thoracicsomites, right side; c, last abdominal somite, telson, and uropod; d, uropod, ventral view,enlarged; e, submedian teeth of telson, ventral view, enlarged.
riorly, inner denticle appressed to margin of submedian tooth, apexat level of intermediate tooth; ventral surface of telson with longi-tudinal carinae, including a short postanal keel, 2 sharp, long carinaeon each submedian tooth, and 1 shorter, less distinct carina on eachintermediate tooth.Proximal segment of uropod with 2 sharp, dorsal tubercles mesial todorsal carina; uropod setation reduced; most of inner margin ofproximal segment of exopod smooth, a few small proximal setaepresent; proximal segment of exopod with fixed ventrodistal spinule
48 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124
overhanging articulation of distal segment; distal segment of exopodsmooth mesially, distolateral setae present; endopod slender, sinuous,most of inner margin smooth, lacking setae, 4-5 small, proximalsetae present; distolateral margins of endopod with normal comple-ment of setae; inner spine of basal prolongation slenderer and shorterthan outer.Color.?Traces of dark chromatophores present across posteriorthird of carapace, sixth thoracic somite, and first, third, fourth, andfifth abdominal somites, pattern most visible on first abdominalsomite; telson with dark spots along anterior margin.Size.?Female, TL 26.7. Other measurements: carapace length5.2; fifth abdominal somite width 4.5; telson length 3.1, width 3.5.Discussion.?G. bicarinatus closely resembles G. demanii Hender-son and G. hendersoni Manning in the reduction of setae on the innerportions of the uropod but differs from both of these and other speciesof the genus in having two ventral carinae on each submedian toothas well as a short, median postanal keel. As in G. demanii, a few proxi-mal setae are present on the inner margin of the endopod in thenew species. The carinate submedian teeth of the telson and theslender uropodal endopod will distinguish this species from the fol-lowing new species, G. crosnieri.Etymology.?The name is from the Latin, "hi", meaning "two,"and "carina", meaning "keel," referring to the bicarinate ventralsurface of the submedian teeth of the telson.Distribution.?Known only from the type-locality, Nosy Be,Madagascar. Gonodactylus crosnieri, new speciesFigure 15Holotype.? 1 9, 17.8; Banc de Pracel, western coast of Mad-agascar; 50 m; muddy sand; A. Crosnier; June 1959; USNM 124098.Paratypes.? 1 9, 19.3; Lagon de Mayotte Id., Comoro Islands;50 m; A. Crosnier; August 1959; USNM 124099. 1 9, 13.5; same;MNHNP.Description.?Anterior margins of rostral plate straight or withslight posterolateral slope; anterolateral angles acute but rounded.Ocular scales small, erect.Mandibular palp and 5 epipods present.Lateral processes of sixth and seventh thoracic somites subtruncate,subequal in size or that of sixth somite slightly larger.Carinae of sixth abdominal somite not markedly inflated, each withapical spiuule.Telson as long as broad, appearing elongate, with 2 pairs of slender,sharp, marginal teeth; lateral teeth not developed, lateral carinafusing with margin of intermediate tooth; dorsal surface of telson
STOMATOPOD CRUSTACEA?MANNING 49
with numerous spinules and tubercles; median carina flask shaped,tapering distally, with apical spine and subapical tubercles; 1-2 rowsof spinules flank median carina, converging posteriorly; anteriorsubmedian carinae with distal tubercle or spinule, flanked laterallyand distally by spinules ; anterior surface of telson with sharp tubercleon each side at level of intermediate carinae of sixth somite; carinaeof marginal teeth each ornamented dorsally with spinules; numeroussmall submedian denticles and 2 sharp intermediate denticles, latterrecessed anteriorly, present; ventral surface lacking distinct longi-tudinal carinae.
Figure IS.
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Gonodactylus crosnieri, new species, female holotype, TL 17.8, Banc de Pracel:a, last abdominal somite, telson and uropod; b, uropod, ventral view.Uropod abnormal in shape and setation; inner margin of exopodsmooth, lacking setae, apex and outer margin of exopod setose;proximal segment of exopod with fixed ventrodistal spinule over-hanging articulation of distal segment; endopod very broad, almosthalf or more than half as broad as telson, inner and outer marginsconvex, lacking setae, distolateral setae present; endopod with long,dorsal carina; spines of basal prolongation subequal in length.Color.?Faded.Size.?Females only known, TL 13.5-19.3. Other measurements offemale holotype, TL 17.8: carapace length 3.6; fifth abdominal somitewidth 2.8; telson length 2.4, width 2.5.Discussion.?G. crosnieri, new species, resembles G. demaniiHenderson, G. hendersoni Manning, and G. bicarinatus, new species,in dorsal armature of the telson but can be distinguished from thesespecies by the absence of lateral teeth on the telson and by the broaduropodal endopod, which is half or more than half as broad as the telson273-102?67 4
50 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124and which has smooth lateral margins and a distolateral fringe ofsetae. In the other three species the outer margin of the narrowendopod is fringed completely with setae. The new species lacks thelongitudinal ventral carinae of the telson and the acute anterolateralangles of the rostral plate that are characteristic of G. hendersoni.The broad uropodal endopod of this species will separate it from allother species in the genus.Etymology.?The species is named for Alain Crosnier, whocollected the species and made available for study most of the othermaterial reported herein.Distribution.?Known only from Madagascar and the ComoroIslands, in 50 m. Gonodactylus demanii Henderson, 1393Gonodactylus demani.? Gravier, 1935, p. 360, fig. 3.Gonodactylus De Manx.?Dollfus, 1938, p. 213 [part].Gonodactylus demanii.?Manning, 1967b, p. 8, fig. 3 [older references].Illustration.?Manning, 1967b, fig. 3.Previous records.?Fort Dauphin (Gravier, 1935); Maromandia(Dollfus, 1938).Material.? 1 broken cf, CL 2.5; Tulear, Madagascar; from off-shore reef; K. Boss; Anton Bruun Cruise 7, IIOE; USNM.Diagnosis.?Anterolateral angles of rostral plate rounded orangled; ocular scales small; telson with 3 pairs of well-developedmarginal teeth, submedians with movable apices; dorsal surface oftelson with numerous spinules and tubercles; inner margin of uropodalexopod smooth, lacking setae; most of inner margin of uropodal endo-pod smooth, 1-5 proximal setae present; submedian teeth of telsoneach with 1 ventral carina.Color.?Faded.Discussion.?As shown by Manning (1967b), G. demanii is thewestern Indian Ocean counterpart of G. hendersoni Manning; itdiffers from the latter in having more rounded anterolateral angleson the rostral plate, 1-5 proximal setae on the uropodal endopod,and in lacking sharp ventral carinae on both the submedian and inter-mediate marginal teeth of the telson. Gonodactylus crosnieri, describedabove, differs from both G. demanii and G. hendersoni in having abroad, inflated uropodal endopod, with most of the inner and outermargins lacking setae but a few distal setae present.Distribution.-?-Western Indian Ocean, from scattered localitiesbetween southern India, the Red Sea, and northern Mozambique(Manning, 1967b).
no. 364i STOMATOPOD CRUSTACEA?MANNING 51Gonodactylus lanchesteri Manning, 1967fGonodactylus De Mani var. spinosus.?Dollfus, 1938, p. 215, fig. 17.Gonodactylus lanchesteri Manning, 1967b, p. 11, fig. 4 [older references].Illustration.?Manning, 1967b, fig. 4.Previous records.?Comoro Islands (Manning, 1967b) ; ?Mada-gascar (Dollfus, 1938).Material.? 1 d\ 25.6; Mayotte Id., Comoro Islands; intertidalzone; A. Crosnier; September 1959; USNM. 1 d\ 27.8; 1 9, 29.9Nosy Be, Madagascar; in sandstone; A. Crosnier; 20 May 1958USNM. 1 9, 15.5; same; A. Crosnier; 10 December 1958; MNHNP1 9, 24.0; same; intertidal zone; A. Crosnier; February 1962; MNHNP1 9, 25.4; same; from hard coral; A. G. Humes; 10 September 1964USNM. 1 o\ 27.3; Ambatoloka, Nosy Be, Madagascar; MmeChavane; July 1958; USNM.Diagnosis.?Rostral plate rounded anterolaterally; ocular scalessmall; telson with 3 pairs of marginal teeth, submedians with movableapices; dorsal surface of telson with numerous spinules and tubercles;uropod with normal setation, margins of endopod, inner portion ofproximal segment, and distal segment of exopod fringed with setae.Color.?Faded in most specimens; in the male from the ComoroIslands there are median and lateral dark patches on the sixth thoracicsomites, a dark patch on the first abdominal somite, traces of a darkband on each abdominal somite, and the ventral surface of the thoraxand the copulatory tubes are dark.Discussion.?The uropod provided with a normal fringe of setaewill immediately distinguish this species and G. spinosus Bigelowfrom G. demanii Henderson, G. hendersoni Manning, and G. crosnieriManning. Gonodactylus hendersoni differs from G. spinosus in havingthe lateral teeth of the telson well developed, projecting, and inhaving the intermediate denticles recessed anteriorly.All of the specimens reported herein have few, large tubercles onthe telson.Distribution.?Western Indian Ocean from the Red Sea south-ward to South Africa (Manning, 1967b).Gonodactylus segregatus Lanchester, 1903FlGUKE 16Gonodactylus chiragra var. segregatus a+ b Lanchester, 1903, p. 448, pi. 23 (figs.6, 7, 7a).Previous records.?None.Material.? 1 9, 22.9; Banc Vert, Madagascar; 14 m; A. Crosnier;January 1959; USNM. 1 9, 17.8; Mayotte Id., Comoro Islands;intertidal zone; A. Crosnier; September 1959; MNHNP.
52 PROCEEDINGS OF THE NATIONAL MUSEUMDescription.?Rostral plate with anterior margins transverse;anterolateral angles acute but not spiniform.Ocular scales small, erect.Mandibular palp and 5 epipods present.Lateral process of sixth thoracic somite more truncate and largerthan that of seventh somite.Carinae of sixth abdominal somite almost tubular, not greatly-inflated, each with apical spinule.
/ I h
Figure 16.
?
Gonodactylus segregatus Lanchester, female, TL 22.9, Banc Vert: a, outline oflateral processes of sixth, seventh, and eighth thoracic somites, right side; b, last abdominalsomite, telson, and uropod; c, uropod, ventral view.Telson as broad as or broader than long, with 2 pairs of sharpmarginal teeth, laterals distinct but not well developed, submedianswith movable apices; dorsal surface of telson lacking tubercles andspinules except for apical armature of dorsal carinae; median carinatapering distally, with posterior spinule; accessory medians short,well developed, with posterior spinule, subapical tubercle present or
no. 3641 STOMATOPOD CRUSTACEA?MANNING 53
absent; accessory medians not fusing with median to form anchor;anterior submedians slender, with apical tubercle; an irregular U-shaped ridge, open anteriorly, extends posteriorly from midlengthof telson between anterior submedian and intermediate carinae,across central carinae, to equivalent point on other side of telson;carinae of marginal teeth sharp, submedians divergent anteriorly;numerous small submedian denticles present; 2 intermediate denticlespresent, each sharp, anteriorly recessed; ventral surface of telsonlacking sharp longitudinal carinae.Endopod and expopod of uropod with normal complement of setae,some distal setae of endopod erect; endopod slender, tapered distally,with long dorsal carinae; proximal segment of exopod with slender,ventrodistal spine at articulation of distal segment; inner spine ofbasal prolongation shorter than outer.Color.?Body light, with traces of darker pigment on dorsum;carapace, sixth thoracic somite, and first abdominal somite each withdark dorsal patch; sixth thoracic somite dark laterally; merus of clawwith dark, dorsal patch.Size.?Females only examined, TL 17.8-22.9. Lanchester's speci-mens were all small, TL 7.5-17.5. Other measurements of female,TL 22.9: carapace length 4.4; fifth abdominal somite width 3.1;telson length 2.8, width 2.8.Discussion.?These two specimens correspond very closely withthe account of the species given by Lanchester (1903); the telsonornamentation most closely resembles that recorded by Lanchesteras usegregatus b."Borradaile (1907) considers this species to be identical with G.affinis de Man, 1902, from Ternate, a form described by de Man as avariety of G. chiragra. Identification of G. segregatus with G. affinismust await direct comparison of type material or adequate redescrip-tion of both species based on type material.The specimens recorded herein are similar to G. choprai Manning,1967b, described from specimens taken in the northwestern IndianOcean. Gondoactylus choprai, however, differs from all other speciesin the genus in lacking a fixed distoventral spine on the proximalsegment of the uropodal exopod.Distribution.?Central and western Indian Ocean, from theComoro Islands, Madagascar, and from several localities in theMaldive Islands (Lanchester, 1903); shallow water to 35 fms.Protosquilla Brooks, 1886The genus Protosquilla includes the species assigned by Kemp(1913) to Gonodactylus Group III. Serene (1952, 1954) commented on
54 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124the affinities of some of the species and listed the species known atthat time.Three species occur off Madagascar.Protosquilla lenzl (Holthuis, 1941)Gonodactylus glaber.?Kemp, 1913, p. 182, pi. 10 (fig. 121).?Serene, 1947, p. 385,fig. 1, pi. 2.Gonodactylus lenzi Holthuis, 1941, p. 288 [new name for G. glaber; older refer-ences].?Tiwari and Biswas, 1952, p. 362.?Serene, 1954, pp. 6, 7, 10,34, 52, 73, figs. 5-7, Ha-c, 13-12.?Manning, 1962, p. 11.?Ingle, 1963,p. 31, fig. 31.Illustration.?Serene, 1947, fig. 1, pi. 2.Previous record.?None.Material.? 1 9, 25.5; Nosy B?; intertidal zone; Macnae; Decem-ber 1958; USNM. 5 9, 25.8-30.0; same; sandstone; A. Crosnier;November 1961; MNHNP. 1 d\ 25.9; Point Fievre, Nosy Be;13?24'21" S, 48?18'33" E; J. Rudloe field no. Jr. 13; IIOE; 26 De-cember 1963; USNM. 1 9, 32.3; same; 13?24.4' S, 48?17.8' E; inter-tidal mud flat, 50 yds. right of Centre d'Oceanographie pier; J. Rudloefield no. 24A; IIOE; 4 January 1964; USNM. 1 9, 26.5; AmbatombokaCrater Point, Nosy Be; 13?24'17" S, 48?13'31" E; from base ofvolcanic rocks; J. Rudloe field no. JR-23; IIOE; 3 January 1964;USNM.Remarks.?These specimens agree well with Kemp's account ofthe species. There is some variation in the amount of pitting on thedorsal bosses of the telson; Serene (1947) also commented on thisvariation.The body is covered with light brown chromatophores aggregatedin bands; darker bands or spots are present on the anterior andposterior third of the carapace and on the sixth and seventh thoracicand first abdominal somites. There is a characteristic dark spotlaterally on the sixth thoracic somite.Distribution.?Indo-West Pacific, from scattered localities be-tween the eastern Indian Ocean and the Philippines. Shallow water.
Protosquilla pulchella (Miers, 1880)Gonodactylus pulchellus.?Kemp, 1913, p. 177, pi. 10 (figs. 117-118).?Holthuis,1941, p. 288, fig. 9b [older references].?Serene, 1954, p. 52 [listed].?Stephen-son and McNeill, 1955, p. 252.?Ingle, 1963, p. 30, figs. 29, 49.Illustration.?Kemp, 1913, pi. 10 (figs. 117, 118).Previous record.?None.Material.?2 d\ 18.6-30.0; 1 9, 28.0; Nosy B6; in sandstone;A. Crosnier; 20 May 1958; USNM. 1 d\ 14.6; same; intertidal zone;A. Crosnier; September 1958; MNHNP. 1 d\ 29.5; same; R. Legendre;
no. 3641 STOMATOPOD CRUSTACEA?MANNING 55July 1959; USNM. 1 9, 30.4; same; A. Crosnier; January 1962;USNM. 1 9, 31.4; same; February 1962; MNHNP.Remarks.?These specimens agree well with Kemp's (1913)account of the species. All specimens have small posterolateral spineson the carinae or bosses of the sixth abdominal somite. The darkpatches on the sixth thoracic somites may be well marked on themales but not on the females.Distribution.?Indo-West Pacific, from scattered localities be-tween East Africa, the Red Sea, and Australia. Shallow water.Protosquilla spinosissima (Pfeffer, 1888)Gonodactylus spinosissimus.?Kemp, 1913, p. 191, pi. 10 (figs. 124-125).
?
Holthuis, 1941, p. 292, fig. 9c [with synonymy].?Serene, 1953, p. 507; 1954,pp. 6, 52 [listed].?Ingle, 1963, p. 31, fig. 30.Illustration.?Kemp, 1913, pi. 10 (figs. 124-125).Previous record.?None.Material.?1 9, 12.8; Nosy Be; intertidal zone; December 1958;MacNae; USNM.Description.?Cornea flattened, faintly bilobed, set obliquely onstalk; ocular scales acute laterally; rostral plate trispinous, medianspine longer than anterolaterals, extending to cornea, anterolateralspines extending just past base of eyestalk; anterior margin of carapacelateral to rostral plate slightly concave, angled anterolaterally;dactylus of claw with basal notch on outer margin; mandibular palp2-segmented; first 4 abdominal somites smooth, margin of fourthangled posterolaterally; fifth somite smooth dorsally, with 3-4longitudinal lateral carinae, posterolateral margin spined; sixthsomite with anterior row of spinules, dorsal surface with 6 irregulargroups of erect spines, each situated on low dorsal bosses indistinctlyseparated by shallow furrows; telson longer than broad, completelyfused with sixth somite, ornamented dorsally with 3 bosses, medianround, situated anteriorly, submedians oval, situated at and posteriorto apex of median cleft; surface of telson covered with erect, slenderspines, arising primarily from dorsal bosses and lateral margins;lateral spines arranged in 3 rows, outermost directed laterally, with9-11 spines; posterior margin of telson with deep median cleft, linedwith spines, and shallower submedian clefts; basal segment of uropodwith 1-2 proximal dorsal spines; outer spines of basal prolongationof uropod longer than inner.Color.?Body mottled brown; carapace and merus of claw withlighter transverse band; telson and uropods lighter than body, lackingdefinite pattern in preservative.Discussion.?This small specimen agrees in all details with theexcellent account of the species given by Kemp (1913).
56 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124ProtosquiUa guerinii (White) is the only other species of Protosquttlaknown from the western Indian Ocean in which the telson is orna-mented with long dorsal spines. That species, however, differs inhaving the fifth abdominal somite armed with 3-5 transverse rows ofspinules, a row of spinules on the uropodal endopod, and the dorsalspines of the telson arranged in an entirely different pattern. Thecentral spines on the telson of P. guerinii form a corona; the threefissures of the telson margin are almost equally deep and are linedwith spinules. In addition, the dorsal spines on the telson of P. gueriniimay be provided with fleshy tips that are absent in P. spinosissima.Distribution.?Indo-West Pacific, from scattered localities be-tween the western Indian Ocean and Japan. It has not been recordedpreviously from Madagascar. It usually is recorded from shallowwater, on reefs, but Tattersall (1906) reported material from 45-50fms off Ceylon. Literature CitedBalss, H.1910. Ostasiatische Stomatopoden. Abh. Math.-phys. Klasse Bayer Akad.Wiss. Miinchen, suppl. 2, trans. 2, pp. 1-11, figs. 1-2.Barnard, K. H.1926. Report on a collection of Crustacea from Portugese East Africa.Trans. Roy. Soc. South Africa, vol. 13, pt. 2, pp. 119-129, pi. 10.1950. Descriptive list of South African stomatopod Crustacea (mantisshrimps). Ann. South African Mus., vol. 38, pp. 838-864, figs. 1-4.1955. Additions to the fauna-list of South African Crustacea and Pycno-gonida. Ann. South African Mus., vol. 43, pp. 1-107, figs. 1-53.1962. New records of marine Crustacea from the East Africa region.Crustaceana, vol. 3, no. 3, pp. 239-245, figs. 1-3.Bigelow, R. P.1894. Report on the Crustacea of the Order Stomatopoda collected bythe steamer "Albatross" between 1885 and 1891 and on otherspecimens in the U.S. National Museum. Proc. U.S. Nat. Mus.,vol. 17, pp. 489-550, figs. 1-28, pis. 20-22.1931. Stomatopoda of the southern and eastern Pacific Ocean and theHawaiian Islands. Bull. Mus. Comp. Zool. Harvard, vol. 72,no. 4, pp. 105-191, figs. 1-10, pis. 1-2.BORRADAILE, L. A.1898. On some crustaceans from the South Pacific: Stomatopoda, pt. i.Proc. Zool. Soc. London, 1898, pp. 32-38, pis. 5-6.1899. On the Stomatopoda and Macrura brought by Dr. Willey from theSouth Seas. In Willey, Zoological results based on material fromNew Britian, New Guinea, Loyalty Islands and elsewhere, col-lected during the years 1895, 1896, and 1897, vol. 4, pp. 395-428,pis. 36-39.1907. Stomatopoda from the western Indian Ocean. In The Percy SladenTrust Expedition to the Indian Ocean in 1905, under the leader-ship of Mr. J. Stanley Gardiner. Trans. Linn. Soc. London, ser.2, vol. 12, pp. 209-216, pi. 22.
no. 3641 STOMATOPOD CRUSTACEA?MANNING 57Bouvier, E. L.1914. Sur la faune carcinologique de l'ile Maurice. C. R. Acad. Sci.,vol. 159, pp. 698-704.1915. Decapodes marcheurs et stomatopodes recueillis a l'ile Maurice parM. Paul Carrie. Bull. Sci. France et Beige, ser. 7, vol. 48, no.3, pp. 178-318, figs. 1-42.Brooks, W. K.1886. Report on the Stomatopoda collected by H. M. S. "Challenger"during the years 1873-76. In Reports on the scientific researchof the "Challenger," Zoology, vol. 16, pp. 1-116, pis. 1-16.Chhapgar, B. F., and Sane, S. R.1967. Two new species of Squilla (Stomatopoda) from Bombay. Crusta-ceana, vol. 12, pt. 1, pp. 1-8, figs. 1-2.Chopra, B.1939. Stomatopoda. In The John Murray Expedition, Scientific Reports,vol. 6, no. 3, pp. 137-181, figs. 1-13.Crosnier, A.1965. Les crevettes penaeides du plateau continental Malgache. CahiersO.R.S.T.O.M., Oceanographie, vol. 3, no. 3, suppl., pp. 1-158,figs. 1-14, maps.Dawydoff, C.1952. Contribution a l'etude des invert^bres de la faune marine benthiquede l'lndochine. Biol. Bull. France Belgique, suppl. 37, pp. 1-158.Dollfus, R. Ph.1938. Stomatopoda, n: Catalogue synonymique des especes jusqu'a pre-sent r?colt6es dans la Mer Rouge, y compris la partie sud duCanal de Suez et le Golfe d'Aden. In Mission Robert Ph. Dollfusen Egypte, x. Mem. Inst. Egypte, vol. 37, pp. 185-236, figs.1-8, 10-11, 13-23.FOURMANOIR, P.1952. Observations sur la faune marine et le peche a l'ile Europa. Mem.Inst. Sci. Madagascar, ser. a, vol. 7, no. 2, pp. 167-188, 1 fig.,pis. 9-13.1953. Note sur le developpement postembryonaire de deux squilles (stoma-topodes) de Madagascar. Nat. Malgache, vol. 5, no. 2, pp. 153-158, figs. 1-13.Gravier, C.1920. Sur une collection de crustaces recueillis a Madagascar par M. lelieutenant DeCary. Bull. Mus. Hist. Nat. Paris, vol. 26, pp.376-383.1935. Crustacds stomatopodes recueillis par M. R. Decary, a Fort-Dauphin(Madagascar). Arch. Mus. Hist. Nat. Paris, ser. 6, vol. 12, pp.357-361, figs. 1-3.1937. Stomatopodes de cotes d'Indo-Chine. Ann. Inst. Oceanogr., vol. 17,fasc. 3, pp. 175-211, figs. 1-23.1938. Stomatopoda, i. In Mission Robert Ph. Dollfus en Egypte, ix.Mem. Inst. Egypte, vol. 37, pp. 163-183, figs, a-d, 1-5.Haan, W. de1833-1850. Crustacea. In de Siebold, Fauna Japonica sive descriptioanimalium, quae in itinere per Japoniam, jusse et auspiciis superi-orum, qui summum in India Batavia Imperium tenent, suscepto,annis 1823-1830 collegit, notis observationibus et adumbrationibusillustravit, pp. i-xvi, i-xxxi, vii-xvii, 1-243, pis. a-q, 1-55, circ. 2.
58 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 124Hansen, II. J.1926. The Stomatopoda of the Siboga Expedition. In Siboga Expedition,Monogr. 35, pp. 1-48, pis. 1-2.Hoffmann, C. K.1874. Crustaces et echinodermes de Madagascar et de l'ile de la Reunion,pp. 1-36, pis. 1-10. In Pollen and van Dam, R6cherches sur lafaune de Madagascar et des ses dependances, vol. 5, pt. 2, pp. 1-58,pis. 1-10.1874a. Enumeration des crustace's trouves a Madagascar et les Mascareignes,pp. 37-44. In Pollen and van Dam, R6cherches sur la faune deMadagascar et des ses dependances, vol. 5, pt. 2, pp. 1-58, pis.1-10.HoLTHTJIS, L. B.1941. The Stomatopoda of the "Snellius" Expedition. No. xn in Biologicalresults of the "Snellius" Expedition. Temminckia, vol. 6, pp.241-294, figs. 1-9.1951. Note on Caledoniella montrouzieri Souverbie, a gastropod molluscliving commensally on stomatopod Crustacea. Basteria, vol. 15,nos. 3-4, pp. 69-71.1964. Preliminary note on two new genera of Stomatopoda. Crustaceana,vol. 7, no. 2, pp. 140-141.Humes, A. G.1965. New species of Hemicyclops (Copepoda, Cyclopoida) from Mada-gascar. Bull. Mus. Comp. Zool. Harvard, vol. 134, no. 6, pp.159-259, pis. 1-36.Ingle, R. W.1963. Crustacea Stomatopoda from the Red Sea and the Gulf of Aden. No.26 in Contributions to the knowledge of the Red Sea. Bull. SeaFish. Res. Sta., Haifa, no. 33, pp. 1-69, figs. 1-73.JURICH, B.1904. Die Stomatopoden der deutschen Tiefsee-Expedition. In Wissen-schaftliche Ergebnisse Deutschen Tiefsee-Expedition, "Valdivia,"vol. 7, pp. 361-408, pis. 25-30.Kemp, S.1913. An account of the Crustacea Stomatopoda of the Indo-Pacific region,based on the collection in the Indian Museum. Mem. IndianMus., vol. 4, pp. 1-217, 10 text-figs., pis. 1-10.Kurian, C. V.1954. Contributions to the study of the crustacean fauna of Travancore.Bull. Centr. Res. Inst., Univ. Travancore, Trivandrum, ser. c,vol. 3, no. 1, pp. 69-91, 3 figs.Lanchester, W. F.1901. On the Crustacea collected during the "Skeat Expedition" to theMalay peninsula, together with a note on the genus Actaeopsis.Proc. Zool. Soc. London, 1901, pp. 534-574, pis. 33-34.1903. Marine crustaceans, vm: Stomatopoda, with an account of the vari-eties of Gonodactylus chiragra. In Gardiner, The fauna and geog-raphy of the Maldive and Laccadive Archipelagoes: Being anaccount of the work carried on and of the collections made by anexpedition during the years 1899 and 1900, vol. 1, pp. 444-459,pi. 23.
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Lenz, H.1905. Ostafrikanische Dekapoden und Stomatopoden gesammelt von HerrnProf. Dr. A. Voeltzkow. Vol. in in Voeltzkow, WissenschaftlicheErgebnisse der Reisen in Madagaskar und Ostafrica in den Jahren1889-95. Abhandl. Senckenb. Naturf. Ges., vol. 27, pp. 341-392,pis. 47-48.1910. Crustaceen von Madagaskar, Ostafrika und Ceylon. In Voeltzkow,Reise in Ostafrika in den Jahren 1903-1905 mit Mitteln der Her-man und Elise geb. Heckmann Wentzel-Stiftung ausgefiihrt, vol.2, pp. 539-576.Lenz, H., and Richters, F.1881. Beitrag zur Krustaceenfauna von Madagascar. Abhandl. benck.Naturf. Ges. Frankfurt, vol. 12, pp. 421-428.Man, J. G., de li>L .1898. Bericht iiber die von Herrn Schiffscapitan Storm zu Atieh, an denwestlichen Kiisten von Malakka, Borneo und Celebes sowie in derJava-See gesammelten Decapoden und Stomatopoden. Sechster(Schluss-) Theil. Zool. Jahrb. Syst., vol. 10, pp. 677-708, pis. 28-38.1929. On a collection of decapod and stomatopod Crustacea from PulauBerhala, an islet situated in the Straits of Malacca. Bijd. Dierk.,vol. 26, pp. 1-26, pis. 1-3.1929a. Decapoda and Stomatopoda from Pulau Berhala. Misc. Zool. Su-matrana, no. 36, pp. 1-3.Manning, Raymond B.1962. Stomatopod Crustacea collected by the Yale Seychelles Expedition,1957-1958. Postilla, no. 68, pp. 1-15, figs. 1-2.1965. Stomatopoda from the collection of His Majesty The Emperor ofJapan. Crustaceana, vol. 9, pt. 3, pp. 249-262, figs. 1-2, pis. 11-12.1966. Notes on some Australian and New Zealand stomatopod Crustacea,with an account of the species collected by the Fisheries Investi-gation Ship Endeavour. Rec. Australian Mus., vol. 27, no. 4, pp.79-137, figs. 1-10.1967. Stomatopoda in the Vanderbilt Marine Museum. Crustaceana, vol.12, pt. 1, pp. 102-106.1967a. A review of the genus Odontodadylus (Crustacea: Stomatopoda).Proc. U.S. Nat. Mus., vol. 123, no. 3606, pp. 1-35, figs. 1-8, pi. 1.1967b. Notes on the demanii section of genus Gonodactylus Berthold withdescriptions of three new species (Crustacea, Stomatopoda). Proc.U.S. Nat. Mus., vol. 123, no. 3618, pp. 1-27, figs. 1-8.1967c. A revision of the family Squillidae (Crustacea, Stomatopoda), withthe description of eight new genera. Bull. Mar. Sci., in press.Miers, E. J.1880. On the Squillidae. Ann. Mag. Nat. Hist., ser. 5, vol. 5, pp. 1-30,108-127, pis. 1-3.1884. Crustacea. In Report of the zoological collections made in theIndo-Pacific Ocean during the voyage of H.M.S. "Alert," 1881-82, pp. 178-322, 513-575, pis. 18-35, 46-52.Monod, Th.1925. Sur les stomatopodes de la cote occidentale d'Afrique. Bull. SocSci. Nat. Maroc, vol. 5, no. 3, pp. 86-93, pis. xx-xxi.
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