Adult and Larval Stomatopod Crustaceans Occurring in Hawaiian Waters1

SIDNEY JOSEPH TOWNSLEy2

INTRODUCTION t~on of the Bernice P. Bishop Museum, found Sl X genera and eight species from Hawaii: THE STOMATOPODA constitute the only order belonging to the division Hoplocarida of the Squilla oratoria, S. alba, Pseudosquilla ciliata, malacostracan Crustacea. All living forms are P. oculata, Coronida sinuosa, Lysiosquilla macu­ ~do~todactylus included in a single family, the Squillidae, lata,. hanseni, and Gonodactylus Since th at time no extensive studies whose characteristics are the same as those guerlnJ. of the order. The order, according to Bigelow have been made on specimens from Hawaii, (1894), may be defined as Crustacea which but occasional reports of previously recorded ~pecies fr~m have these characteristics : the stalked eyes and this region have found their way anrennules are borne upon distinct mo vable into .the ht.erature. In addition to these eight segments; the adult rostrum is separated from speCles which have been again collected dur­ the carapace by a movable joint; the carapace _ ing the progress of thi s study, I have found is small and does not cover the last four tho­ two species which have not been recorded fr~.m racic somites; the first five pairs of thoracic this region previousl y, namely, Squilla appendages are not biramous and serve as boops and an undescribed species of Squilla. accessory mouth parts, with the second pair Stomatopods are usually found in very shal­ strongly developed into large raptorial limbs ; low water, but some species occur at greater the last three pairs of thoracic appendages depths. I have collected Gonodactylus guerini are adapted for walkin g, are biramous and bear in Hawaii only at depths over 100 meters, and a lateral segment upo n the penultimate seg­ Kemp (1913) reports th at Squilla leptosquilla ment; the abdomen is well developed; fila­ was dredged by the "Investigator" at a depth mentous gills are carried upon the exopod of of 370-419 fathoms . The adults are bottom the first five abdominal appendages; the sixth dwellers where the y may burrow into the sub­ ~nh ab i~ abd ominal appendages, the uropod s, with the strate .or crevices in coral. They are telson, act as a powerful tail fan. seclusive In their habits and durin g the day At the present time the six recognized gen­ are usually found either in their burrows or era have a world -wide distribution , bein g but a short distance away. At times they wan­ found in the tropical , subtropical, and rem­ der some distance from the ir burrows, as is perate waters of the Atlantic, Pacific, and In­ shown by the fact that the y are occasionally dian oceans. All six genera are represented in captured in crab nets and at submersible lights the Hawaiian fauna. Edm ondson (1921), re­ used for collecting at night. Inasmuch as the porting upon the stomatopods in the collec- burrows are quite deep (a foot or more below the surface), and because the animals retire 1 C ontr ibuti o~ No, 37, Hawaii Marine Laboratory, Prep ar e ~ as partial !.equirement for the M .S. degree. to the bottom of them at the least alarm University of H awaii. Manuscript received November stornatopods are .quite difficult to collect ; 14, 1951. 2 Graduate assistant, Osborn Zoological Laborator y consequently, the gro up has been studied Yale University. ' very little, the most notable works being those

399 400 PACIFIC SCIENCE, Vol. VII, October, 1953 of Miers (1880), Brooks (1886), Bigelow withstanding their great abundance, only two .(1894), and Kemp (1913). Of these works, or three early larval forms have been traced the most useful for the identification of the to their adult counterpart. Their long plank­ Hawaiian species has been Kemp's mono­ tonic existence easily accounts for the world­ graph on the Indo-Pacific species. wide distribution of the genera and the pres­ Stomatopods are carnivorous and readily ence of species such as Pseudosquilla ciliata seize any flesh held in front of their burrows . in the Pacific Ocean , Indian Ocean, Red Sea, This behavior has provided the most satis­ and Atlantic Ocean . At present there appear factory means for capturing them , as they can to be onl y two species which are endemic in be lured from their burrows by suspending Hawaii, namely, Coronida sinuosa and Squilla fish flesh a short distance away and captured calumnia n. sp. when they attack the flesh. Considerable dex­ Attempts were made to rear larval spec­ terity is necessary, however , as they withdraw imens, collected in plankton tows, in the lab­ rapidly into their burrows . It has been im­ oratory, but this proved to be impos sible as possible to keep more than one specimen at their period of survival in captivity was very a time in small aquaria as they are highly short, and none of the larvae ever molted cannibalistic. while in an aquarium . Because rearing of the It is doubtful whether the adults often serve larvae was unsuccessful, it was necessary to as food for other animals, although they may study specimens collected in plankton tows occasionally be found in the stomach con ­ and at night light stations and material from tents of fish; bur the planktonic larval stages the stomach contents of pelagic fishes. which compose a substantial proportion of The earlier authors such as Miers (1880), the neritic plankton form a considerable part in their work on larval sromatopods, were ofthe diet of pelagic fishes. content with bestowing separate generic and Instead of carrying their incubating eggs trivial names upon the larvae. If the adult attached to the abdominal appen dages, as do genus to which the larval specimens belonged most Malacostraca, the stomatopods deposit was known, they added the suffix -erichthus them at the bottom ofthe burrows where they to the first syllable of the adult genus plus are aerated by currents of water produced by a trivial designation .(except in the case of the abdominal appendages of the parent. alima of Squilla). Thus Pseuderichthus communis These appendages are paddle shaped and con­ Hansen, according to this system of classifi­ form to the cylindrical shape of the hole . cation, represented the larval form of a species The eggs perish when they are deprived ofthis of Pseudosquilla. Because no keys to these constant current of water, and, as the female individual larval species were included, future will not remain with the eggs when placed authors found it difficult to identify specimens in an aquarium, no larvae have been reared in their possession with those which had been from the egg in captivity. previously described , and new larval species The young do not hatch as nauplius larvae. were erected to compensate for this. Claus All known forms hatch in an advanced stage (1872) and Brooks (1886) have done the most which bears many adult characteristics , but notable works on the larval forms . Giesbrecht it is difficult to assign the early larvae properly (1910), Foxon (1932), and Gurney (1942) because the diagnostic characters do not ap­ have reviewed the works of Claus and Brooks pear until later. and expanded their results to include other The larvae are delicate, transparent organ­ unidentified larvae. However, I have found isms which lead a planktonic existence. The that the material has not been adequately larval life is long, and they pass through many simplified below the level of the genus, be­ molts before reaching the adult stage. Not- cause in many cases they have retained the Hawaiian Stornatopods - TOWNSLEY 401 previously assigned generic and rrivial names The present study was undertaken because of rhe larvae, thus making it difficult to as­ recent investigations on the feeding hab its of certain the relationship of these species to pelagic fishes have shown that the larval stages those collected in Hawaii. Most of the larvae of the Stornatopoda 'rank second in impor­ belonging to a particular genus resemble one tance as food for certain species of tu na in another so closely that it is virtually impossi­ the Hawaiian area, and that they are signifi­ ble to place them in their proper specific cant in the diet of several other pelagic fishes. category without rearing them to maturity. Inasmuch as the adult stages of the pelagic Because of these difficulties, I have not tried larvae involved are found on the reefs and to relate the Hawaiian larval forms with those shores of the Islands , it is apparent that the ofthe previously described species, but where general economy of the neritic waters in the it has been possible I have attempted to cor­ vicinit y is influenced to some extent by the relate them with their proper adult species. productivity ofthe reef and shore fauna. These The specific identification of the larvae was two points suggest a third, namely, that pe­ made in the manner used by Claus (1872) lagic fishes may be influenced to venture close and Brooks (1886) in which three methods to shore because of the increased food supply were emplo yed: (1) a comparison of larval there. stages with posdarval forms which show char­ In order to assess objectively the impor­ acteristics of both the larvae and adults, in tance of these larval stages in the general this manner working from the adult back economy of the neritic realm, it is necessary through the larval series to the earliest iden­ to be able to identify the species involved, tifiable stage; this is by far the most accurate so that a rapid qualitative andquantitative means of identification, except, of course, analysis ofboth the plankton and the stomach where rearing of the larvae to the adult has. contents of pelagic fishes may be made. The been accomplished; (2) a comparison of the present problem was designed to accomplish distribution of larval forms and known adults these ends for the Stomatopoda, and it is in which it is possible to make specific diag­ hoped that the results obtained will prove of noses with some accuracy in those cases where use to workers in other fields. only one or two adult species are known to A cknowledgments occur, but cases may arise in which the adult specimen is unknown; (3) the relative abund­ I wish to express my thanks to Dr. C. H . ance of larval and adult species, although this Edmondson forpermitting me to examine the is the least accurate method used, has proved specimens in the Bernice P. Bishop Museum, to be useful when one species is known to be to Mr. Vernon Brock for making available more numerous than others and it is assumed specimens collected by the Territorial Divi­ that the larval forms also retain the same sionof Fish and Game, and to the staff of the relative numerical relationship. Pacific Oceanic Fishery Investigations for pro­ Almost all authors who have worked with viding specimens collected by them . I am larval stornatopods have noted that invariably indebted to Dr. A. H. Banner for his constant there are forms present which in no respects help and criticism on the collection of mate­ bear any resemblance to the adults recorded rial and preparation of the manuscript, and for the region. This is also true ofthe material to Dr. R. W. Hiatt for suggesting the problem at hand from Hawaii. This would seem to and for criticism of the manuscript . In addi­ indicate that there are numerous species tion I would like to extend my thanks to Mr. throughout the world , probably from greater Kenji Ego , Mr. Daniel Yamashita, and Miss depths, which are known only in their larval Winifred Tseu for their help in collecting stages. speCImens. 402 PACIFIC SCIENCE, Vol. VII, October, 1953

ADU LT HAW~IIAN SQUI LLIDAE the species of each natural group resemble each other far more closely than they do those The genera of Squillidae easily fall into two of the other group. groups which are distinguished by differences Squilla is separated from the other members in the ischio-rneral articulation of the raptorial of the first group in that its species all possess claw (second maxilliped). In one group the longitudinal carinae on the body segments . articulation between the merus and ischium Pseudosquilla, Lysiosquilla, and Coronida are sep­ is terminal, whereas in the other the ischium arated from one anoth er by the shape of the articulates at a point anterior to the proximal bod y, the carapace, and the telson , as is in­ end of the merus. This character is the basis dicated in the following key and descriptions. for the primary division of the family, with Odontodactylus and Gonodactylus are readily dis­ Squilla, Pseudosquilla, Lysiosquilla, and Coronida tinguished because species of Odontodactylus comp rising the first group, and with Gono­ possess teeth on the inner margin of the dac­ dactylus and Odontodactylus comprising the lat­ tylus of the rapto rial claw, whereas species of ter group. Although the phylogenetic sig­ Gonodactylus lack such teeth . nificance of the ischio-rneral articulation of To avoid confus ion in terminology, Figure the raptorial claw is not clearly understood, 1 presents characteristics used in the key and

A A A B B B C 0 C C E 0 F E 0 G F H G I E H L I b v J d K A VI B V C VII L VI 0 VIII A E a M F C N G H 0 I J E i< a L c F

FIG. 1. Diagrams illustrating morph ological details and terminology of sroma topods. a, Rostru m, carapace, exposed th oracic somites, and first abd ominal somite of Squilla. A-I , Structures of carapace. A, Rostrum; s, anterolateral spi ne; c, median carina; D, gas tric groove; E, intermediate carina; F, lateral carina; G, marginal carina; H, cervical groove; I, pos teriorly reflexed marginal carina. J-N, Structures of rhoracic and abdo minal somires. J, Anterolateral spi ne of fifth rhoracic somite; K , submedian carina; L, intermediate carina; M, lateral carina; N , marginal carina. V-VIII, Fifth to eighth thoracic somites. I, First abdominal somite. b, Secon d th oracic append age (raptorial claw). A, Carp us; B, meru s; c, propo dus ; D, dact ylus ; E, ischium. c, Tels on and last two abdominal somites of Squilla. A-E, Structures of abdomina l sornites. A, Submedian carina; B, later al carina; c, margin al carina; D, intermediate carina; E, med ian carin a. F-L, Srructures of telson . F, Pre­ lateral denticle; G, lateral "to oth ; H, lateral denticle; I, intermediate to ot h; J, intermediate denticles; K, sub­ med ian denticles; L, submedian tooth. V-VI, Fifth and sixth abdominal sorn ires. d, Last abdominal somi te and telson of Odontodactyias. A-D, Structur es of sixth abdominal som ite (VI).A, Sub. medi an carina; B, lateral carina; c, second intermediate carina; D, first intermediate carina. E- L, Structures of telson . E, Marginal carina ; F, seco nd lateral carina; G, interme diate carina; H, sub median carina; I, median carina; J, lateral tooth; K, intermediate to oth ; L, submedian tooth. e, Lateral aspect of head and carapace of Squilla. A, Cornea of eye; B, mandibular palp ; c, anrennal scale; D, firsr thoracic appendag e; E, third th oracic appendage; F, seco nd th oracic appendage (raptorial claw). Hawaiian Sromatopods-TOWNSLEY 403

in the descriptio ns of the species . Th ese have length from tip of rostrum to tip of been partially adapted from Kemp (1913), submedian spines of telson . whose termin olog y h as been em plo yed ...... Squilla oratoria throughout this work . DD. Dactylus of raptorial claw with 5 teeth ; body not reachin g over 90 mm . in Key to Adult Hawaiian Sguillidae length from tip of rostrum to tip of A. Raptorial claw with articulatio n of me­ submed ian spines of telson . rus and ischium termin al (Fig. 3g), me­ ...... : " Squilla boops rus grooved ventrally for reception of E. Submedian teeth on posterior margin of propodus throughout its length, pro­ telson with mo vable tips ; 5 or 6 mov­ podus finely pectinate or with a series able spines along outer margin of exo­ of fixed spines along the upper margin, pod of uropod ; 2 rounded lobes be­ dactylus not inflated at base B tween inner and outer spine of basal AA. Raptorial claw with ischio-meral arti­ prolongation of uropod; body opague culation at a point anterior to the prox­ white throughout Squilla alba imal end of the merus (Fig. 21c), ventral EE. Submedian teeth on posterior margin surface of merus grooved for reception of telson without movable tips; 8 or 9 of propodus for not more than 0.75 of mo vable spines alon g outer margin of its length, propodus may or may not be exopod of uropod ; 1 rounded lobe finely pectinate along the upper margin , near base of inner spine of basal pro­ dactylus inflated at base (O donrodac. longation of uropod ; body with definite tylus and Gonodactylus) I brown pigment markin g carinae and B. Carapace with well-marked carinae .cer­ margins Squilla calumnia n. sp. vical groove defined across th e dorsal F. Lateral margins of exposed thoracic surface; first 5 abdominal somites with somites and abdomi nal somites parallel longitudinal carinae (Squilla) C throu ghout, not increasing in breadth BB. Carapace witho ut well-marked carinae, posteriorly; telson with sharp median ' cervical groove not extending over dor­ carina and with lateral carinae (Pseu- sal surface; first 5 abdominal somites dosquilla) G witho ut longitudinal carinae F FF. Lateral margins of expo sed thoracic and C. Anterolateral angles of carapace armed abdominal somites not parallel throu gh ­ with short, acute spines; cornea of eyes out, abdominal somites increasing in set obliguely on stalk (Fig: 3a), nearly breadth posteriorly; tels on with median 1.5 times as wide as greatest breadth carina low and rounded, and with out of stalk, distinctly bilobed D lateral carinae (Lysiosquilla and Cor- Cc. Anterolateral angles of carapace round­ onida) H ed, without short acute spines (occa­ G . Inner spine of basal prolongation of sionally with spines in S. alba); cornea urop od lon ger than outer spine; eyes of eyes set transversely on eyestalk (Fig . small and cylindrical; 6 carinae besides 6), slightly wider th an the greatest median carina on telson . breadth of stalk, very slightly bilob ed ...... Pseudosquilla ciliata ...... E GG. Inner spine of basal prolongatio n of D . Dactylus of raptorial claw with 6 teeth ;" uropod shorter than outer spine; eyes bo dy usually more than 100 mm . in flattened, club-shaped; 8 carinae be- - - - sides median carina on telson . 3 The number given.for th e t.eeth on rhe raptori al dacryl1;1s t h ro u g~ ou t this study includes the termi nal ...... '.' .. . Pseudosquilla oculata tooth In all species, except Odontodactylus hanseni. H. Alternating b ands oflight and dark pig- 404 PACIFIC SCIENCE, Vol. VII, October, 1953

ment throughout length ofbody; rapto­ toward gastric grooves. Rostrum longer than rial dactylus with 9 or 10 teeth o~ inner wide, subquadrate, with small median spine, margin; telson with low, rounded, me­ lateral margins convergent anteriorly, not dian carina . . . .Lysiosquilla rnaculata coverin g ophthalmic somite.Eyes large ; cor­ HH. No alternating bands of light and dark nea bilobed, set transversely on stalk; anterior pigment throughout length of body; margin ofophthalmic somite slightly truncate raptorial dactylus with 4 teeth on inner between base of eyestalks (Fig. 3a). Merus margin; telson closely studded with of raptorial claw articulating terminally with large tubercles in definite patterns .. ... isc hium , ventral surface longitudinally · Coronida sinuosa grooved throughout its length for reception I. Dactylus of raptorial claw with 9 teeth of propodus; propodus finely pectinate along on inner margin; telson with ordinary uppe r margin and with four sharp movable longitudinal carinae; rostrum with even­ spines; dactylus not inflated at base, with six ly convex anterior border and evenly long, sharp teeth on inner margin. Propodus rounded angles ; eyes large, breadth of of first, third, and fourth thoracic appendages cornea about equal to 0.5 the length of nearly circular, that of fifth subcircular, being the carapace; preserved specimens yel- slightly longer than wide (Fig. 3[ , h, i ). lowish white or pink . Mandibular palp of th ree segments (Fig. 3b, · Odontodactylus hanseni c). Free thoracic and abdominal somites dor­ II. Dactylus of raptorial claw without teeth soventrally flattened, all possessing longitu­ on inner margin; telson with concentric dinal carinae (first five abdominal somites rings of flesh-tipped spines; rostrum trispinous ; eyes small, breadth ofcornea about equal to 0.12 5 the length of the carapace; preserved specimens mottled red or reddish brown . · Gonodactylus guerini

Squilla oratoria de Haan Figs. 2, 3a-k Squil/a aratoria de Haan, 1844: pl. 51, fig. 2 . [figures only]; 1849: 223 [description]. Squil/a affinisBerthold, 1845: 26, pl. 3,figs.1- 2. Squilla nepa Miers, 1880 : 25. Chloridella affinis de Man, 1907 : 439 .

DES CRIPTION: Carapace consid erably nar­ rower anteriorly than posteriorly; nearly twice aslon g (including rostrum) as greatest breadth anteriorly; conspicuous gastric and cervical grooves , the latter continuous across mid­ dorsal region; seven distinct, longitudinal ca­ rinae, median bifurcated anteriorly for about 0.25 the distance to cervical groove, inter­ mediate extending nearly entire length of ca­ rapace,marginal producedanteriorly into sharp spine and posteriorl y rounded and reflexed FIG. 2. Squilla aratoria de Haan (male). Hawaiian Stomatopods- TOWNSLEY 405

... .. e

•& o

s

f

IOmm

FI G. 3. Squilla oratoriade Haan. a, Anterior segments of adult ; b, right pars molaris of mandible and mandibular palp (ventral aspect); c, right pars molaris of mandible (dorsal aspect); d, right pars incisiva of mandible; e, right maxilla; f, g, right first and secon d rhoracic appendag es; b, right thir d and fourth th oracic appendages; i, right fifth th oracic appendag e; j, telson and urop ods ; k, right first abdo minal app endage of male. each with eight carinae); fourth through dominal somite with six spines, two sub­ eighth thoracic somites exposed; intermediate median, two intermediate, two lateral. Telson carinae and lateral carinae of abdominal som­ with a single median crest, with radiatin g ites 1-5 end in short, sharp spines; sixth ab- series ofpits on each side ; distal margin armed 406 PACIFIC SCIENCE, Vol. VII, October, 1953

with three pairs of large immovable teeth, DISTRIBUTION: This species appears to be submedian, intermediate, and lateral; three to somewhat more restricted in distribution than five submedian denticles, seven to nine inter­ are other stomatopods, being reported only mediate denticles, and a single lateral denticle, from the Hawaiian Islands,Guam, Philip­ also a pair of prelateral denticles. Exopod of pine Islands, inland waters ofJapan, and the uropods with nine movable spines on outer China Sea. margin. Basal portion of uropod elongated into two spines, inner longer than outer, a Squilla hoops Kemp small lobe near the outer edge of the inner Figs. 4, 5a-f spine (Fig. 3j ). Accessory reproductive organ of male shown in Figure 3k. The dorsal sur­ Squilla boops Kemp, 1911 [May]: 97. face of the specimens is somewhat pitted and Squilla quadraticauda Fukuda, 1911 [August]: a dull mottled tan in color, the ventral surface 287, pI. 11, figs. 3-5. is highly polished and somewhat lighter in DESCRIPTION: Carapace broad anteriorly, color. Specimens vary from 145 to 200 mm. breadth ' behind anterolateral spines exceeds in length from the anterior end ofthe rostrum 0.5 the total length including the rostrum; to the extremity of the submedian spines of conspicuous gastric and cervical grooves, the the telson. latter continuous across the middorsal region; DISCUSSION: Several varieties of this species seven distinct longitudinal carinae, median are recognized, but the Hawaiian specimens discontinuous anterior to cervical groove, end­ seem to fit the characteristics given by Kemp ' ing in a small pit, marginal produced anterior­ (1913) for those specimens which he had ex­ ly into a short spine and posteriorly rounded amined from this region. He found that spec­ and reflexed toward gastric grooves. Rostrum imens from Hawaii did not seem to fit any of broader than long, lateral margins upturned, the described varieties and did not attempt to rounded and slightly convergent anteriorly, place them in any definite variety . a small median carina. Ophthalmic somite Three males and three females collected at exposed, anterior margin truncate between Oahu were examined. All were collected in eyestalks. Eyes large; cornea distinctly bi­ regions with a muddy bottom suitable for lobed, set obliquely on stalks, considerably burrowing. The Bishop Museum has eight more than 0.3 length of carapace; prominent specimens, two females collected at Guam, lobe on external aspect of eyestalk. Isch io­ and two males and four females bought at a meral articulation of raptorial claw terminal; Honolulu market. All were very similar in ventral surface of merus longitudinally appearance, except for small variations in col­ grooved throughout its length for reception or and size; characteristics which do not seem of propodus; carpus with three carinae; pro­ to be of systematic importance. There seems podus finely pectinate along upper margin to be no apparent sexual dimorphism. 'The ' and with two movable spines proximally; only positive means found for distinguishing dactylus not inflated at base, with a small males from females was the penis at the base ventral tubercle, five teeth on inner margin of the eighth thoracic appendage, and the ac­ (Fig. 5b). Propodus of third and fourth thor­ cessory reproductive organ of the male found acic appendages rounded posteriorly, those on the first abdominal appendage (Fig. 3k). of first and fifth nearly straight and parallel This species is commonly taken in crab nets to anterior margin (Fig. 5a, c-e ). Mandibular by fishermen at the Ala Wai Canal, Honolulu. palp composed of three'segments, similar to Occasionally they appear in the Honolulu fish that of S. oratoria (Fig. 3b). Free thoracic and markets, their large size making them desir­ abdominal somites dorsoventrally depressed, able for food. increasing in breadth posteriorly; all possess- Hawaiian Stomatopods- TOWNSLEY 407 ing longitudinal carinae; fourth through eighth thoracic somites exposed, lateral mar­ gin of fifth produced into an anteriorly di­ rected spine and a rounded posterior lobe, lateral margin of sixth of two rounded lobes, seventh and eight rectangular; first five ab­ dominal somites with eight carinae, the mar­ ginal and lateral ending in a short acute spine, all intermediates except that ofthe first somite ending in spines, submedian of fifth ending in spines ; sixth abdominal somite with six carinae, submedian, intermediate, and lateral all ending in spines . Telson with a single median crest terminating in a short spine; radiating series of pits on each side ofmedian crest; distal margin armed with three pairs of immovable teeth , submedian, intermediate, and lateral; three to five submedian denticles , seven to nine intermediate denticles, one lat­ eral denticle, also a pair ofprelateral denticles. Exopod ofuropods with seven movable spines along outer margin . Basal portion of uropods elongated into two spines, inner longer than

FIG. 5. Squilla boops Kemp. a,b,c,d,e, Right first, second, third, fourth, fifth thoracic appendages; f, right first abdominal appendage of male.

outer, a small rounded lobe between the two (Fig. 4). Accessory reproductive organ of male shown in Figure Sf Kemp mentions that the type had the median portion of the carapace, abdomen, and telson covered with small, gray chromatophores. These were not sufficiently abundant to detract from the gen­ eral coloration, which was yellowish in his type specimen but faintly pink in the spec­ FIG. 4. Squilla boops Kemp (male). imens at hand . The posterior half of the fifth 408 PACIFIC SCIENCE, Vol. VII, October, 1953

abdominal somite between the lateral and DESCRIPTION: Carapace longer than great­ marginal carinae and the outer angles of the est breadth, slightly narrower anteriorly than sixth somite are black in all three of these posteriorly, with distinct gastric and cervical specimens, as well as in Kemp's type spec­ grooves, with five longitudinal carinae, an­ imen. No mention of the color of Fukuda's terolateral angles rounded and usu ally armed spe cimen was made. The th ree local spec ­ with short acute spines, posterolateral angles imens vary from 68 to 75 mm.; Kemp's was bro adly rounded. Rostrum longer than broad, 89 mm., and Fukuda's 40 mm. truncate or triangular. Ophthalmic somite par­ DISCUSSION: One male and two female spec­ tially exposed, anterior margin straight be­ imens of this species have been examined. tween base of eyestalks, with small rounded They were taken from the stomach contents process at base of each eye dorsally. Eyes of black skipjack, Euthynmtsyaito Kishinouye, . large ; cornea set transversely on stalks, only caught off Moku M anu [Island] off the north­ slightly broader than greatest breadth ofstalk, east coast of Oahu. One specimen, a male, about 0.3 total length of carapace, including was but slightly digested, so the characteris­ rostrum. Antennular peduncle about same tics given by Kemp for this species could be length as carapace, including rostrum. Ischio­ easily recognized. meral articulation of raptorial claw terminal ; S. boops can be readily distinguished from ventral surface of merus grooved longitudin­ S. aratoria by its smaller size and by the pres­ ally through out its length for reception of ence of only five teeth on the raptorial dac­ propodus; carpu s with a blunt median carina; tylus and only seven spines on the outer outer margin of propodus finely pectinate , margin of the uropod. Early in thi s study it th ree movable spines near proximal end (only was th ou ght that these were small specimens one shown in Fig . 7b, other two not visible of S. aratoria, but the differences noted abo ve in this view); dactylus not inflated at base, have shown them to be the ad ults ofKemp's notched inferiorly, outer margin straight or Squil/a boops. slightly con cave, inner margin armed with Kemp and Fuku da's specimens were both six teeth. Anterior and posterior margins of females; thus previou sly no description has third and fourth th oracic appendages round­ been made of the modified accessory repro­ ed, first and fifth straight and nearly parallel ductive orga n of the male found on the first (Fig. 7a, c- e) -" . No mandibular palp. Fifth abdominal appendage. However, Kemp's through eighth th oracic somites expos ed ; lat­ (1913) figures of the female have been useful eral ma rgin of fifth seen in do rsal view com­ in identifying the present specimens, which posed of two distinct processes, a long, an­ I consider conspecific. . teriorly directed spine and a rounded posterior DISTRIBUTION: The type specimen, a female, lobe; sixth, seventh,.and eighth not bilobed, was taken by the " Investigator " in the Gulf but rounded. Lon gitudinal carinae on all ex­ ofMartaban, Burma, at a depth of 67 fathoms. posed somites, eight on first five abdominal Another female was collected at Matsuwa, somites, none ending in spines ; posterolateral Sagami Province, Japan. One male and two margins acute and rounded ; sixth abd ominal females have been obtained from the sto ma ch somite with six carinae ending in spines, two . contents of a black skipjack (Euthynnus yaito) submedian, two intermediate, two lateral. Tel­ caught off M oku Manu (Bird Island), Oahu. son with indi stinct median carina but a small median spine posteriorly; radiating series of Squilla alba Bigelow pits on dorsal surface; armed with six mar­ Figs. 6, 7a-f ginal teeth , two submedian with movable tips, Squil/aalba Bigelow,1893: 103;1894:539-541, two intermediate, two lateral; five or six min­ pI. 22. [Complete description and figures.] ute submedian spinules, 11 or 12 intermediate, Hawaiian Stomatopods- TOWNSLEY 409 one lateral. Exopo d of uropods with five or SIX movable spines along their outer margin. Basal portion of uropods elongated into two spines, inner longer than outer, with a dis­ tinct, rounded lobe near base of each spine (Fig. 6). First abdominal appendage shown in Figure 7f Body of specimens an opaque white, eyestalks yellow with a very black cornea. Abdominal and thoracic somites and · carapace with large, white pigment cells dis­ tribute d at random over their surface. Telson , seen from above, nearly transparent so that tissue projecting into it can be easily seen and produces the pattern shown in Figure 6. Spec­ imens range in size from 20 to 47 mm . in length. DISCUSSION : A small female specimen, 20 mm. long, was collected by Mr. Ken ji Ego

FIG. 7. Squilla alba Bigelow. a,b,c,d,e, Right first, second, th ird, fourt h, fifth tho racic app endages; f, right first abdominal appe ndage of fem ale.

at Kawaihae, Hawaii, in a night light station. This is a very striking species, as the color of the specimens immediately attracts atten ­ tion. Its shape is also somewhat peculiar, the carapace and exposed portion of the thorax being equal in length and together making up nearly 0.44 ofthe total length of the bod y. The entire surface of the body is smooth and polished, interrupted occasionally by distinct carinae. The lateral margins of the fifth thor­ acic somite with their long, anteriorl y directed spines and rounded posterior lobes are also FIG. 6. Squilla alba Bigel ow (female). characteristic of this species. 410 , PACIFIC SCIENCE, Vol. VII, October, 1953

The type specimen was collected by Bige­ waihae specimen is considerabl y smaller, be­ low at Bimini Harbor, Bahamas. Edmondson ing only 20 mm . in length. (1921) reported that a female specimen was DISTRIBUTION: The type specimen was col­ taken on Waikiki Reef, Oahu. Unfortunately, lected in calcareous sand at Bimini Harbor, I have not been able to locate this specimen Bahamas. In the Hawaiian Islands one spec­ in the Bishop Museum for comparison with imen was taken from dead coral on Waikiki the specimen taken at Kawaihae. Edmondson Reef, Oahu, and another at Kawaihae, Hawaii, states that the specimen which he collected at a night light. These are the only records agreed with Bigelow's previous description known for this species. for the species. The Kawaihae specimen agrees with Bigelow 's description in color, shape of SquiOa calumnia n. sp. the eyes, general body proportions, shape of .Figs. 8, 9a-f the lateral margin of the fifth thoracic somite, shape of the telson, and the presence of two DESCRIPTION: Carapace, excludin g rostrum, lobes between the inner and outer spines of about as long as greatest breadth; slightly the uropods. However, there are certain char­ narrower anteriorly than posteriorly; distinct acteristics in which the specimens differ con­ gastric grooves and cervical groove; median, siderably. Bigelow's type possesses definite intermediate, lateral, and marginal longitu­ longitudinal carinae on the carapace, the ros­ dinal carinae present, median discontinuous trum is truncate, the anterolateral angles of and bifurcated anterior to cervical groove, the carapace are rounded and armed with a lateral posteriorly reflexed toward gastric short spine , and the outer margin of the uro­ groove; anterolateral and posterolateral angles pods is armed with six movable spines. The unarmed and rounded. Rostrum broader than Kawaihae specimen, on the other hand , does long, truncate, small median carina, lateral not have any carinae on the carapace, the margins upturned and rounded anteriorly. rostrum is distinctly triangular, the antero­ Ophthalmic somite exposed, anterior margin lateral angles of the carapace are rounded and concave between base of eyestalks. Eyes large; unarmed, and the outer margin-of the uropods cornea set transversely on stalks, only slightly is armed with only five movable spines. I broader than greatest breadth of stalk, about believe that these differences are due ro the rel­ 0.25 total length of carapace, including ros­ atively immature condition of the Kawaihae trum. Antennular peduncle about same length specimen , and that it is probably the last as carapace, 'excluding rostrum. Ischio-meral postlarval stage of this species. It is possible articulation of raptori al claw terminal; ventral that the subsequent molt would reconcile the surface of merus longitudinally grooved differences between this and the Bimini spec­ throughout its length for reception of pro­ imen. podus; carpus with median and lateral carinae; propodus finely pectinate along outer surface, Bigelow makes little note ofthe submedian three movable spines near proximal end ; dac­ spines on the posterior margin of the telson. tylus not inflated at base, outer margin slight­ His figure (1894: pi. 22) is indistinct, but ly concave, armed with six teeth. Anterior there appears to be a line indicating that they margin of propodus of third , fourth , and possess movable tips. In the Kawaihae spec­ fifth thoracic appendages straight (Fig. 9c-e), imen the tips of these spines are unquestion­ posterior margins rounded; propodus of first ably movable. thoracic appendage nearly circular (Fig. 9a). No male specimens have been reported. Mandibular palp composed of two segments. The largest of Bigelow's specimens was 41 Fifth through eighth thoracic somites ex­ mm. , Edmondson's was 47 mm. , and the Ka- posed, with longitudinal carinae; lateral mar- Hawaiian Stornatopods - TOWNSLEY 411 gin of fifth, seen in dorsal view, composed of two distinct processes ,anteriorly directed spine and nearly straight posteriorspine; lateral mar­ gin of sixth nearly straight, angular; seventh rounded posteriorly. Eight longitudinal cari­ nae on first five abdominal sornites, subme­ dian, intermediate, lateral, marginal; all mar­ ginal carinae terminating in a short acute spine; sixth abdominal somite with six cari­ nae, all ending in spines, two submedian, two intermediate, two lateral. Telson with median carina, ending in a small median spine pos­ teriorly; radiating series of pits on each side

FIG. 9. Squilla calumnia n. sp. a,b,c,d,e, Right first, second, third, fourth, fifth thoracic appendages; f, right first abdominal appendage of female.

of median carina; armed with six teeth, two submedian, two intermediate, two lateral; four submedian denticles, seven intermediate denticles, one lateral denticle. Outer margin of exopod of uropods with eight movable spines on one side, nine on the other. Basal portion ofuropods elongated into two spines, FIG. 8. Squilla calumnia n. sp. (female). inner longer than outer, distinct rounded 412 PACIFIC SCIENCE, Vol. VII, October, 1953 lobe near base of inner. First abdominal ap­ rounded posterior lobe . Here both processes pendage shown in Figure 9[ The coloration are distinctly angular, the anterior spine of the specimen is sufficient to distinguish it curved forward, the posterior one projec ting from others which I have examined. The laterally. The shape of the sixth thoracic som­ ground color is an opaque white, which is ite is distin ctive in this species as well, in partially obscured by dark-bro wn chromato­ that the others have rounded lateral margins phores along the posterior bor der of the thor­ or are distin ctly bilobed. acic and abdominal somites and which also TYPE SPECIMEN: U.S.N .M . 93097. clearly mark all the lon gitudinal carinae of the body . The cornea of the eyes is a deep Pseudosquilla ciliata (Fabricius) black, the eyestalks retaining the opaque white of the body. Specimen 26 mm. lon g. Figs. 10, lla-m TYPE LOCALITY: A single male specimen : Squilla ciliata Fabricius, 1787: 333. 26 mm. long was collecte d by the Territorial Cancer ciliatus Linnaeus, 1790: 2990. Division of Fish and Game at a night light Cancer (mantis) [sic] ciliatus Herbst, 1796: 102. station at Hilo Dock, Hilo, Hawaii. Squilla stylifera Lamarck, 1818: 189. DISCUSSION: Alth ough the penis which is Squilla quadrispinosa Eydo ux and Souleyet, located at the base of the eighth thoracic 1841: 262, pl. 5, fig. 1. appendage is present, the specimen is ap­ Pseudosqaill« stylifera Dana, 1852: 622, pl. 12, parently immature, as the accessory repro ­ figs. 4a-e. ducti ve organ on the endopod of the first abdomi nal appendage is undeveloped. In DESCRIPTION: Carapace longer than great­ spite of this condition I feel that the other est breadth, slightly narrower anteriorly than characteristics are sufficiently distinct for the posteriorly with conspicuous gastric grooves, separation of this specimen from the other cervical groove wholly absent, lacking cari­ species in this .genus. nae, anterolateral and posterolateral angles At first glance this species and Squilla alba unarmed and rounded. Rostrum rounded an­ appear to be very similar; but the immovable teriorly, about as wide as lon g, covering the submedian spines along the outer margin of ophthalmic somite. Eyes tubular, cornea fol­ the telson, the more numerous spines along lowing long axis of stalk (Fig. 11a). Merus the .outer margin of the uropods, the shape of raptorial claw articulating terminally with ofthe lateral margin of the fifth thoracic som ­ ischium, ventral surface grooved throughout ite, and the single lobe at the base of the its length for reception of propodus; propo­ inner spine of the uropod will serve to dis­ dus with fine pectinations along outer margin tinguish this species from S. alba. The round­ and with four movable spines near proximal ed anterolateral angles of the carapace, the end; dactylus with three (sometimes four) large globular eyes, and the six teeth on the lon g, sharp teeth . Propodus of fifth thoracic raptorial dactylus are shared by both these appendage subcircular, slightly longer than species and will serve to separate them from wide (Fig. 11i ). Mandibular palp of three the other members of this genus. segments. Free thoracic and abdominal som­ The fifth thoracic somite in this specimen ites compress ed, all lacking longitudinal cari­ appears to be composed of two distinct proc­ nae; sixth, seventh, and eighth thoracic sorn­ esses when seen from above, but it is different ites exposed; posterolateral angles of abdom­ from others showing this same condition. In inal somites with small, sharp spines; sixth others , e.g., S.laevis Hess (1865), S. hieroglyph­ abdominal somite with four spines, two sub­ ica Kemp (1911), and S. alba Bigelow (1893), median, two intermediate. Telson with seven there is a spine directed anteriorly and a carinae, one median, two submedian, two Hawaiian Stomatopods- TOWNSLEY 413 intermediate, two lateral; armed with six spines, two submedian with movable tips, two intermediate, two lateral; no submedian denticles, two intermediate denticles, single lateral denticle. Exopod of uropods with nine (sometimes ten) movable spines on outer margin . Basal portion of uropod elongated into two spines, inner longer than outer. Ac­ cessory reproductive organ of male shown in Figure 11/. Specimens vary from 35 to 85 mm . in length. DISCUSSION: Fifty-eight specimens of this well-known species were examined , of which 23 were males and 35 were females. All were collected from the reefs around Oahu in water not over 2 fathoms deep. This species repre­ sents the dominant form of the Hawaiian Islands . It has been collected from both Kauai and Hawaii where it seems to be as abundant as it is at Oahu. The species appears to prefer the rough substrate of the large exposed reef flats such as those found in Kaneohe Bay, Oahu. It has not been found in those regions having a substrate of coarse shifting sand or strong wave action . FIG. 10. Pseudosquilla ciliata (Fabricius) (male) . There appears to be no clearly defined sex­ ual dimorphism, and the only definite means to be quite variable, but this variability is that I have found for distinguishing males not sufficient to warrant their being placed from females lies in the penis found at the in any other group. I have examined three base of the eighth thoracic leg (Fig. llj) and specimens from Kaneohe Bay, Oahu, in the accessory reproductive organ of the male which there were ten spines rather than nine found on the first abdominal appendage (Fig. on the exopod of the uropod; and one spec­ 111). Specimens exhibit all degrees ofcoloring imen in which there were nine spines on one from an olive drab through yellow to a light side and ten on the other. Two specimens in tan with bright red, blue, and yellow chroma­ the collection have the outer spine ofthe basal tophores. The dactyli of the thoracic append­ prolongation of the uropod longer than the , ages are, in most cases, light rose. inner. This is a character which has been con­ According to Brooks (1886) the Atlantic sidered to be specific for P. oculata (Brulle), specimens ofthis species differ in certain small but all other characters of this specimen are details from those which are found in the those of P. ciliata. Indo-Pacific region . Borradaile (1899) named Specimens vary in length from 38 to 85 the Atlantic specimens var. occidentalis, but mm. Generally this species is somewhat larger later workers such as Tattersall (1906) and than its congener, P. oculata, and in the field Kemp (1913) have shown that the characters they can usually be distinguished in this way. used by Borradaile are to be found in the DISTRIBUTION: There are 12 species of Indo-Pacific specimens as well.. The speci­ Pseudosquilla, nine of which are found in the mens from the Hawaiian Islands a~so appear Indo-Pacific region . P. ciliata and its Hawaiian 414 PACIFIC SCIENCE, Vol. VII, October, 1953

~r-, ~iI~ ~ j

FIG. 11. Pseadosqailla ciliata (Fabricius). a, Anterior segments with rostrum removed; b, right antenna with scale; c, right pars molaris of mandible and mandibular palp ; d, right pars incisiva of mandible; e, right maxiIJa;f,g, right first and second thoracic appendages;h, right third and fourth thoracic appendages;i, right fifth thoracic appendage; j , eighth thoracic appendage offemale (top) and male showing penis (bottom) ; k,l, right first abdominal append­ ages of female and male; m, tel~on and uropods. Hawaiian Stomatopods- TOWNSLEY 415

relative, P. oculata, are both known from the Pacific, Indian, and Atlantic oceans . Ordina­ rily they are found in the intertidal area in these regio ns, but Bigelow (1891) reports that P. ciliata was collected at a depth of 20- 23 fathoms. P. ciliata has been reported from the Hawaiian Islands, Fiji Islands, So­ lomon Islands, Loyalty Islands , Japan, New Britain , New Guinea, Red Sea, Madagascar, Bermuda, Florida Keys , Puerto Rico , and St. Thomas.

Pseudosquilla o culata (Brulle) Figs. 12, 13 Squilla oculata Brulle, 1836: 18, fig. 3. Squilla monodactyl« A. Milne-Edwards, 1878 : 232. Pseudosquilla oculata Miers, 1880: 1l0, pI. 3, figs. 3, 4. Pseudosquilla mOl1odactyla Miers, 1880: 11 0, pI. FIG. 12. Pseudosquilla oculat« (Brulle ) (male). 3, figs. 1, 2. DESCRIPTION: Carapace lon ger than great­ with two sublateral spines ; sixth abdominal est breadth, slightly narrower anteriorly than somite with two submedian and two sub­ posteriorly, with conspicuous gastric grooves, lateral spines . Telson with nine carinae, one cervical groove wholly absent, lacking carinae, median, two submedian, two intermediate, anterolateral and posterolateral angles un­ two sublateral, two lateral; armed with six armed and rounded. Rostrum with small me­ spines , two submedian with movable tips , dian spine, about as wide as long, coverin g two intermediate, two lateral ; no submedian ophthalmic somite. Eyes flattened, appearing denticles, two intermediate denticles, single somewhat club-shaped rather than tubular lateral denticle. Exopod of uropods with ten (compare Figs. lla and 12), cornea set trans­ (sometimes ll) movable spines along outer versely on stalk. Merus of raptorial claw ar­ margin.Elongated basal portion of uropod ticulating terminally with ischium, ventral with outer spine lon ger th an inner (Fig. 12). surface grooved longitudinally throughout its Accessory reproductive organ of male shown length for reception of propodus; propodus in Figure 13. All specimens were a mottled with fine pectinations along upper surface and tan and all possessed a dark spot on the lateral with four movable spines near proximal end ; edges of the carapace, as noted by Kemp dactylus with th ree long, sharp teeth . Propo­ (1913). The dactyli are tipped with rose . Spec­ dus of fifth thoracic appendage subcircular, imens vary from 35 to 70 mm . in length. slightly longer than wide. M andibular palp DISCUSSION: This species superficially re­ of three segments. Free thoracic and abdom­ sembles the more common P. ciliata. In Ha­ inal somites compressed, all lackin g longi­ waiian waters the two species are commonly tudinal carinae; fifth, sixth, seventh, and found in the same habitat, and , except through eighth thoracic somites exposed; postero­ close scrutiny, P. oculata may often be mis­ lateral angles of abdominal somites with taken for P. ciliata. The eyes of this species small, sharp spines ; fifth abdominal somite are very characteristic in shape and contrast 416 PACIFIC SCIENCE, Vol. VII, October, 1953

DISTRIBUTION: This species and P. ciliata are the only members of this genus which have been found in the Hawaiian Islands. As mentioned previously, P. oculata is found in the same environment as P. ciliata. Specimens are recorded from Samoa and Hawaii in the . Pacific, from the Macclesfield Bank in the South China Sea, from Mauritius in the In­ . dian Ocean, and from the Cape Verde Islands and Madeira in the Atlantic.

Lysiosquilla maculata (Fabricius) Figs. 14, 15a- g Squilla arenaria terrestris or locusta Rumphius, 1705: 4, pi. 3, fig. E. Squilla maculata Fabricius, 1793: 51l. Cancer (mantis) [sic] arenarius Herbst, 1796: FIG. 13. Pseudosquilla oculata (Brulle), right first ab- 96, pi. 33, fig. 2. dominal appendage of male. . Squilla arenaria Randall, 1839: 146. well with those of P. ciliata (d . Figs. lla and DESCRIPTION: Carapace nearly as broad as . 12). The spine-tipped rostrum, the extra pair long, anterior breadth equal to posterior of carinae on the telson, and the longer outer breadth. .smooth, strongly convex from side spine of the uropod will all serve to place to side, gastric grooves distinct, cervical specimens in this species. groove obsolete or wholly absent, no longi­ This species is common in Hawaii: the tudinal carinae, anterolateral and postero­ Bishop Museum collection has five males and lateral angles bro adly rounded. Rostrum six females, and in addition I have examined broader than long, smooth, lateral margins one male and four females. The largest spec­ convergent anteriorly to an acute but blunt imen was a male 70 mm. long, and the small­ apex, small longitudinal carina on anterior est was 35 mm. long. All specimens were one third. Ophthalmic somite anteriorly ex­ collected at Oahu in shallow water together posed, dorsally produced into a pair of lobes with specimens of P. ciliata. terminating acutely on each side of rostral Variability in color and form does not seem apex. Eyes large ; cornea set obliquely on to be pronounced in this species. One spec­ stalk, breadth about equal to greatest length imen has 11 spines on the outer margin ofthe of cornea and stalk combined. Antennular exopod of one uropod and 10 on the other. peduncle with small acute dorsal spine, less This variation parallels the condition found than one-half length of carapace, excluding in P. ciliata and indicates that this chracteristic rostrum. Ischio-rneral articulati on of raptorial is quite variable. claw terminal; merus gro o ved ventrally There is no clear sexual dimorphism. The throughout its length for reception of pro­ accessory reproductive organ on the first ab­ podus; carpus with blunt dorsal carina ter­ dominal appendage of the male is shown in minating in a sharp spine which overhangs Figure 13. If it is compared with Figure 11/, anterior margin; propodus finely pectinate on it can be seen that both P. oculata and P. ciliata upper margin, four movable spines near prox­ are very similar with respect to this structure. imal end; dactylus slender, 9 to 11 teeth Possibly this indicates strong affinities. (usually 10), outer margin straight or slightly Hawaiian Stomatopods- TOWNSLEY 417 concave (Fig. 15d). Propodus of third and ' fourth thoracic appendages orbicular with two weak spines at proximal end, those of first and fifth nearly rectangular and lacking spines (Fig. 15c, f). Fourth through eighth thoracic somites exposed, lateral margins rounded, lacking longitudinal carinae. Abdominal so­ mites depressed ;first through fifth lacking lon- ' gitudinal carinae, increasing in breadth pos­ teriorly; sixth narrower than preceding ,so­ mites, with distinct grooves on each side of mid-line. Telson broader than long, convex dorsally, concave ventrally, both sides marked by a feeble median triangular elevation lim- ' ited by a pair of posteriorly convergent

FIG. 15. Lysiosquilla maculata (Fabricius). a, Right pars molaris of mandible and mandibular palp; b, right pars incisiva of mandible; c,d, right first and second thoracic appendages; e, right third and fourth thoracic appendages; f, right fifth thoracic appendage; g, right first abdominal appendage of male. FIG. 14. Lysiosquilla maculata (Fabricius) (female). 418 PACIFIC SCIENCE, Vol. VII, October, 1953

grooves; sometimes marked by patches of 'with those which they report as having these pits rather regularly disposed near lateral mar­ modifications. gins; posterior margin indistinctly notched This species and Squilla oratoria are the two in mid -line with three pairs of blunt lobes, largest stornatopods found in the Hawaiian the two outermost generally sharper and more Islands . I have examined six males and three conspicuous than the inner. Inner dorsal edge females ranging from 70 to 185 mm. from . of peduncular segment of uropods ending in the tip of the rostral spine to the posterior short spine; exopod of uropods with eight border of the telson. Other authors have re­ or nine movable spines; basal portion of uro­ ported specimens ranging from 69 to 283 mm. pod elongated into two long spines, inner Lysiosquilla maculata seems to occur in shal­ nearly twice length of outer, but obscured low water with a sandy or muddy substrate dorsally. Modified first abdominal appendage where it may burrow easily. All those spec­ of male shown in Figure 15g. Animals are imens which I have examined had been col­ marked by alternating series of light and dark lected in the fine silt on the reefs at Oahu and bands throughout the length of the body. Hawaii . Mr. Kenji Ego (personal communi- . Specimens vary in length from 69 ro 283 mm. cation) reports that they occur in abundance DISCUSSION: This species is very readily in the mud flats at Kawaihae, Hawaii. identified. All specimens which have been DISTRIBUTION: This species is widely dis­ hitherto reported, as well as those which I tributed throughout the Indo-Pacific. In the have examined, have possessed the character­ Pacific it has been reported from the Hawaiian istic dark transverse bands shown in Figure Islands, Marquesas Islands, Samoa, Fiji, Phil­ 14. In large female specimens this appears to ippine Islands, and the Dutch East Indies. be a secondary sexual characteristic, inasmuch as the carapace, free thoracic somites , and sixth abdominal somite are almost entirely blue-black. Nearly always in males there are alternating dark and light bands with the tel­ son having a pattern which may vary from that shown in the figure but which usually has dark central and lateral areas. Preserved specimens retain the coloration, but the dark pigment becomes a brownish black. There appear to be other characteristics showing a sexual dimorphism in this species. Miers (1880) and Kemp (1913) record several large females in which the carpus and the basal part of the raptorial propodus bear tufts of long hairs, and the latter with only two movable spines at its proximal end instead of four. The raptorial dactylus in these cases is also reported by them to have only a series of eight or nine denticles rather than the long teeth usually found there; however, none of the specimens which I have examined has shown these modifications. Evidently this is due to the relatively small size of the female specimens which I have examined compared FIG. 16 Co ronida sinuosa Edmondson (female). Hawaiian Stomatopods- TOWNSLEY 419

FIG. 17. Coronida sinuosa Edmondson. a, b, Right first and second th oracic appendages; c, right third and fou rth thoracic appendages; d, right fifth thoracic appendage; e, right first abdo minal appendage of female.

In the Indian Ocean it has been reported from ed.Rostrum small, rounded anteriorly, breadth Penung, Tuticorin, Madras, Mysore, and D ur­ greater than length, not coverin g ophthalmic ban . In the Atlantic Ocean adult specimens somite.Cornea of eyes set transversely on have been collected at Antigua, D utch West ' eyestalks, slightl y bilobed; eyestalks narrow , Indies, and larvae from near the Cape ofGood elongated, flattened. Merus of raptorial claw Hope. articulating terminally with ischiurn (Fig. 17b), venrrals urfacelongitudinall ygroovedthrough­ Coronida sinuosa Edmondson out its length for reception of propodus; pro­ Figs. 16, 17a-e podus finely pectinate along upper margin; dactylus notched and sligh tly inflated at base, Coronida sinuosa Edmondson, 1921: 295, fig. 2. with four sharp teeth on inner margi n. First, DESCRIPTION : Carapace very slightly nar­ third, fourth , and fifth thoracic appendages rower anteriorly than posteriorly, nearly twice shown in Figure 17a, c, d. Mandibular palp as long (including rostrum) as breadth at an­ absent. Free thoracic and abdominal somites terior border, lacking distinct carinae, cervical dorsoventrally depressed, all lacking longi­ and gastric grooves wholly absent, antero­ tudinal carinae; fifth through eighth thoracic lateral and posterolateral angles slightly round- somites exposed, lateral margins of fifth with 420 PACIFIC SCIENCE, Vol. VII, October, 1953

an acute, anteriorl y directed process, sixth, found on the posterior margin of the telson , seventh, and eighth narrowed but evenly and I am inclined to believe that there is but rounded (Fig. 16); dorsal surface of sixth a single pair of submedian spines, the rest abdominal somite ornamented with a scroll­ being small, marginal denticles , although Ed­ like series of tubercles (carinae?), nearly syrn­ mondson (1921) recognized two other pairs. metrical in arrangement. Telson with a series The slightly inflated base of the dactylus of similar tubercles which may be unsym­ suggest,s affi nities to Gonodactylus and Odonto­ metrical except in the median region (Fig. dactylus, but the articulation of the ischium 16); posterior border medially notched giving and merus of the raptorial claw indicates a the appearance of two rounded lobes, with a close relationship to Squilla, . Pseudosquilla, pair of submedian spines and four to six mar­ and Lysiosquilla. ginal spines lateral to each submedian.Exo­ Because of their very small size it can be pod of uropo ds with 8 to 11 movable spines seen why this species has not been collected on outer margin ; elongated basal portion of more often. The three specimens which I have uropods with two spines, inner longer than examined ranged from 9.5 to 17.0 mm . from outer. First abdominal appendage of female the tip of the rostrum to the distal end of the shown in Figure 17e. living animals are trans­ submedian spines of the telson . living adults parent white, preserved specimens opaque are also very transparent and difficult to see, white throughout. Specimens range from 9.5 to 17.0 mm. in length. DISCUSSION: Three female specimens ofthis small species have been examined ,including the type specimen in the Bishop Museum. All had been collected from amon g dead coral on Waikiki Reef, Oahu. The species is apparently closely allied to Coronida multituberculata (Borradaile) (1898), but differs considerabl y in the shape of the rostrum and in the ornamentation of the sixth abdominal somite and telson . Specimens may be easily recognized by the ornamentation ofthe last two body segments. The tubercles (or carinae as they are called by .Edmondson) of the sixth abdominal somite form a definite pattern which is nearly sym­ metrical on each side of the median line, suggesting a single median carina, paired sub ­ median carinae, paired intermediate carinae, paired lateral carinae, and paired marginal ca­ rinae. The telson is similarly ornamented, but is not usually as symmetrical as shown in Figure 16. Here there is also a suggestion of a median carina, paired submedian carinae, and paired intermediate carinae; but all are somewh at joined to one another forming a scroll-like pattern. There appears to be con­ siderable variation in the number of spines FIG. 18. Odontodactylus hanseni (Pocock) (male) . Hawaiian Sromatopods - TOWNSLEY 421

especially so in their native habitat on the reefs. Preserved specimens become opaque white or slightly yellowish, with the cornea of the eyes remaining very dark. It is difficult to say whether or not the specimens are mature and whether there is a distinct sexual dimo rphism , because all spec­ imens are females and none were ever collect­ ed with egg masses. The absence of the gastric grooves and the very elongated antennal so­ mites suggests that possibly they are in an immature stage. Edmondson (1921) states that the specimen in the Bishop Museum was kept alive for a month, during which time it molted with no apparent increase in size or change in structure. DISTRIBUTION : All specimens collected have come from dead coral heads on Waikiki Reef, Oahu.

Odontodactylus hanseni (Pocock)

Figs. 18, 19a-f Gonodactylus hansenii Pocock, 1893: 477, pI. 20, figs. 3, 36. Odontodactylus latirostris Borradaile, 1907: 212, pI. 22, figs. 3, 3a. f DESCRIPTION : Carapace longer than great­ est breadth, slightly narrower anteriorly than posteriorly, with distinct gastric grooves, cer­ vical groove wholly absent, longitudinal cari­ FIG. 19. Odontodactylus hanseni (Pocock). a,b,c,d,e, Right first, second, third, fourth, fifth tho racic append­ nae lacking, anterolateral and posterolateral ages; f, right first abdominal appendage of male. angles unarmed and rounded. Anterior margin of rostrum evenly rounded, twice as broad as fifth thoracic appendages somewhat shield­ long. Ophthalmic somite exposed, dorsal shaped (Fig. 19c-e). Mandibular palp of three process deeply excavate anteriorly . Eyes large, segments. Free thoracic and abdominal so­ globular, corneal diameter 0.25 to 0.5 the mites laterally compressed, distinct longitudi- length of carapace, excluding rostrum (Fig. . nal carinae present only on sixth abdominal 18). Ischio -meral articulation ofraptorial claw somite; sixth, seventh, and eighth thoracic situated at a point in advance ofproximal end somites exposed , lateral borders rounded; pos­ of merus (Fig. 19b); ventral surface of merus terolateral angles offourth and fifth abdominal grooved longitudinally for not more than 0.6 somites ending in a short acute spine ; sixth its length for reception of propodus ; upper abdominal somite with four pairs of longitu­ surface of propodus not pectinate; dactylus dinal carinae, one submedian pair ending in inflated at base, armed with 9-11 teeth on its spines, one intermediate pair without spines, inner margin. Propodus of third, fourth , and one second lateral pair ending in spines, one 422 PACIFIC SCIENCE, Vol. VII, October, 1953 marginal pair ending in spines. Telson with clesfield Bank, South China Sea, at a depth high median carina ending in a short terminal of 35 fathoms . Bigelow (1931) reported four spine, submedian, intermediate, second lat­ males, one female, and two juveniles taken eral, and marginal carinae; armed with three in Hawaiian waters by the "Albatross" in pairs of strong marginal teeth, submedian 1902 at depths ranging from 24 to 83 fathoms. (with movable tips), intermediate, and lateral; The specimens which I have examined are in no submedian denticles but a series ofminute agreement with those previously reported. spinules, two intermediate denticles, and a Some characteristics were relatively difficult single lateral denticle. Exopod of uropods to distinguish, but I attribute this to the fact with 10 or 11 movable spines on outer margin. that the animals were considerably disfigured Basal portion of uropods elongated into two from having been in the stomach. Edmondson spines, inner shorter than outer. First abdom­ (1921) has illustrated the telson ofthis species inal appendage of male shown in Figure 19/ more clearly than is shown in Figure 18. Color of the specimens is a uniform light There appears to be no distinct sexual di­ yellowish pink above with the exposed thor­ morphism. The first abdominal appendage of acic and first four abdominal somites having the male shown in Figure 191 is incomplete a light pink posterior margin . The large glob­ because the fixed limb in all three specimens ular eyes are dark brown and the flagella of was indistinguishable. the antennae are pink. The animals are quite small with the reported range in length from 27 to 80 mm . DISCUSSION: In dorsal aspect this species very closely resembles small specimens of Psuedosquilla, but the ischio-meral articulation of the raptorial claw and the large globular eyes are suitable for distinguishing members of this genus from those of Pseadosquilla. The inflated dactylus and the ischio-meral articula­ tion of the raptorial claw are features which members of this genus have in common with Gonodactyltts, but the numerous carinae on the telson, the nonpectinate upper margin of the raptorial propodus, and the series of teeth on the inner margin of the raptorial dactylus will serve to separate this group from the latter. As stated by Bigelow (1894) this genus seems to occupy an intermediate position between Gonodactyltts and Pseudosqailla. Five female and three male specimens of this species have been examined, all of which were taken from stomachs of the skipjack, Kutstaoonus pelamis, and of the yellowfin tuna, Neotbunnas macropterus. The Bishop Museum has one male and two females which were dredged offWaikiki, Oahu, at a depth ranging from 30 to 50 fathoms . Pocock's (1893) type specimen was a single female from the Mac- FIG. 20. Gonodactylus guerini White (female) . Hawaiian Stomatopods- TOWNSLEY 423

~': a

FIG. 21. Gonodactylus guerini White. a, Right pars molaris of mandible and mandibular palp ; b, c, right first and second thoracic appendages; d, right third and fourth thoracic appendages; e, right fifth thoracic appendage; f, left first abdominal appendage of male (endopod only).

Evidently this species is quite common in rounded, lacking distinct carinae, gastric deeper water because all other specimens have grooves well defined, cervical almost obso ­ been taken at considerable depths. Apparently lete. Rostrum quadrate, trispinous, covering this species forms a considerable part of the ophthalmic somite. Eyes small, tubular; cor­ diet of the tunas, inasmuch as the adults and nea bilobed, set transversely on stalk. Ischio­ larvae are often found in stomach contents. meral articulation of raptorial claw situated This indicates that this species is probably at a point in advance ofproximal end ofmerus semiplankronic, because the tunas feed in so that merus extends backward beyond the mid-water or at the surface. These stornato­ joint (Fig. 21c); ventral surface of merus lon­ pods certainly spend some time on the bot­ gitudinally grooved for not more than 0.75 tom because other specimens have been its entire length for reception of propodus; dredged from a rough substrate. upper margin of propodus with a series of DISTRIBUTION: Specimens have been re­ fine acute teeth; dactylus inflated at base, ported from the Hawaiian Islands, the South elongated into a single acute tooth, inner China Sea, the East Indian Archipelago, and margin of dactylus serrated. First, third, the Indian Ocean . fourth , and fifth thoracic appendages shown in Figure 21b, d, e. Mandibular palp present, Gonodactylus guerini White three segments (Fig. 21a). Free thoracic and Figs. 20, 21a-f abdominal somites laterally compressed, all lacking longitudinal carinae (indication of Gonodactylus guerinii White, 1861a: 43, pl. 7; slight marginal carinae on first five abdominal 1861b: 480. somites) ; fifth through eighth thoracic so­ Protosquilla guerinii Brooks , 1886: 75, pl. 16, mites exposed, sligh tlynarrower than posterior figs. 1, 6. width of carapace; posterior portion of fifth DESCRIPTION: Carapace rectangular, nearly abdominal somite bears several rows of short as wide as long, slightly narrower anteriorly, spines, separated from smooth anterior part anterolateral and posterolateral angles broadly by a curved transverse line; sixth abdominal 424 PACIFIC SCIENCE, Vol. VII, October, 1953 somite armed with numerous long spines, and the last two abdominal somites. The each ending in a blunt, rounded tip from rectangular carapace with its trispinous ros­ which protrudes a soft fleshy process. Arti­ trum is found in several species of this genus. culation between telson and preceding seg­ Gonodactylusandits related genus ,Odontodac­ ment barely discernible dorsally. Telson cov­ tylus, are very similar with regard to the ischio­ ered with series of22 regularly arranged spines meral articulation and the inflated base of the similar ro those oLpreceding segment; distal raptorial dactylus (Fig. 21c); however, Gono­ margin composed of four spine-like processes dactylus is distinguished by the absence of bearing secondary spines on outer edges, two spines on the raptorial dactylus, which may submedian with single row of secondary be entirely smooth or serrated on its inner spines, two intermediate with single inner margin (Fig. 21c). Odontodactylus, on the other row of secondary spines and double row of hand, has several spines on the inner margin secondary spines along outer border (Fig. 20). . of the raptorial dactylus. Exopod ofuropods with eight movable spines Brooks (1886) placed this and several allied on outer margin; endopod with five spines species in a new genus, Protosquil/a, because on dorsal surface; elongated basal portion of the telson and sixth abdominal somite, al­ uropods with two spines, outer longer than though separate, were nearly fused. He also the inner. Endopod of first abdominal ap­ stated that the small size of the antennary pend age of male shown in Figure 21f. Spec­ scales and uropods and the trispinous ros­ imens, when alive, are a brilliant red or orange. trum were sufficiently distinct to warrant the Specimens preserved in alcohol immediately removal of these species from the genus Gono­ turn a mottled yellow or tan, bur those pre­ dactylus.Kemp (1913) and Edmondson (1921), served in formaldeh yde have retained their however, did not consider these differences of red color after several months.Specimens a generic nature and retained the original range in size from 28.5 to nearly 70 mm. from name applied to the species by White (1861). the tip of the rostrum to the distal margin of Two specimens were kept alive for observa­ the telson. tion in an aquarium. Both were females, one DISCUSSION: This is the only species of this of which was collected with a mass of fertil­ world-wide genus which has been reported ized eggs; the other laid a mass of eggs in from the Hawaiian Islands. The type spec­ captivity. Both specimens survived very well imen was taken by H .M.S. "Herald" at Fiji even though they were both dredged from a Islands . Miers (1880) reported this to be a depth of over 100 fathoms. The fertilized female, but Kemp (1913) stated that it was eggs developed until the time ofhatching but a male; apparently Kemp was the one who then began to degenerate. Several eggs from made the error in identification. Another fe­ the mass were removed at intervals and pre­ male was taken at Honolulu by H.M.S. "Chal­ served in 7 per cent formalin solution with lenger," but Brooks (1886) put it in the genus the hope that eventually they might be used Protosquilla. The only other record of the spe­ to describe the embryology of this species. cies is a single male specimen in the Bishop One female molted once while in captivity Museum which was taken offWaikiki, Oahu. and remained a mottled yellow afterward. In the spring of 1949 two males and four Little has been written about the function of females were dredged off Waianae, Oahu, in the elaborate spines on the telson ; however, water 50-120 fathoms deep. This appears to my observations have shown that the animals be the greatest number ofspecimens collected either burrow into the substrate or select holes at anyone time. in dead coral which are just large enough for This species is easily recognized by the them to crawl into . Ordinarily they are found radiating series of spines found on the telson partially emerged from their burrows but when Hawaiian Sromatopods - TOWNSLEY 425

disturbed they turn around and close the and by later development into an erichth us opening with their telson, which thus serves type of larva. The pseudozoea hatches with as an effective protective mechanism. only the first and second thoracic appendages, DISTRIBUTION: This species has been re­ the second being in the form of a raptorial ported from Fiji and Honolulu, from rela­ claw as in the adult, a fully segmented ab­ tively deep water , the only record being 50­ domen bearing four pairs of functional pleo­ 120 fathoms. pods; later these develop into either an erich­ thus- or an alima -type of larva. Species of LARVAL HAWAIIAN SQUILLIDAE Lysiosquilla and Coronida hatch as antizcea, l arval stornatopods form a considerable later developing into erichthus larvae ; species part of the neritic plankton throughout the of Gonodactylus, Odontodactylus, and Pseudo­ warmer waters of the world. They are trans ­ squilla all hatch as pseudozoea and later de­ parent organisms which, at hatching, have velop as erichthus larvae. Species of Squilla many adult characteris tics; however, the diag­ hatch as pseudozoea but later develop an nostic features of the adult do not appear alima-type of larva. until late in development, thu s making it dif­ Foxon (1932), following th e work begun ficult to associate the larval forms with the by Hansen, attempted to bring the classifica­ proper adults . The first attempt to draw up tion of the larvae into agreement with the a key to the larvae was made by Bigelow divisions of the adults as presented by Kemp (1894); however, he based his key on the (1913). In doing this he has set down the general appearance of the forms, a character­ morphological feat ures which serve to identi­ istic which is often very deceptive. H ansen fy the larvae from their earliest stages un til (1895) made a careful study of the larvae and the time they take on th e characters observed divided them into two major gro ups, the in the ad ults. Brooks (1886), Giesbrecht alima-type larvae of Squilla and the erichthus­ (1910), Hansen (1926), and Foxon (1932) type larvae of Lysiosquilla, Coronida, Odonto­ have all noted that the only constant differ­ dactylus, Gonodactylus, and Pseudosquilla, on the ence between alima and erichthus larvae is basis of the shape of the propodus of the the number of intermediate denticles between third, four th , and fifth thoracic appendages. the sub median and the two lateral denticles This division was thought by Hansen to em­ of the telson. In the former there are always phasi ze th e relationship of the various genera more than four ofthese denticles, whereas in to one another, but Kemp (1913) and Foxon the latter there are less than four. If one uses . (1932) do not believe that this is of ph ylo­ these denticles as a diagnostic character, it is genetic importance, because none ofthe larval then possible to divide the larval forms into types possesses characteristics which can be the same major groups as the adu lts . In using considered as an ancestral condition from th is system of classification the larval gro ups which the rest of the genera developed. are the same as those of the adults , the like­ Giesbrecht (1910) has noted that in the ness becoming greater at each successive molt. Stomato poda two types of larvae occur at This method of classification is unques­ hatching, but the differences between them tionably suitable for distinguishing the gen­ gradually disappear through a succession of era to which the larvae belong, but it is a later stages. These two early larval types have much more difficult task to assign them to been designated pseudozoea and antizoea. their proper species. In no species has an The aritizoea is characterized at the time of individual in an early larval stage been reared hatching by biramous appendages on the first to the adult form . In dividuals in later stages five thoracic somites, an abdomen which is have been reared to the postlarval condition, usu ally unsegmented and lackin g pleopods, but they are so delicate that laboratory con - 426 PACIFIC SCIENCE, Vol. VII, October, 1953 ditions must be ideal, requiring water which B. Larvae hatch with raptorial claw (Fig. is well aerated and free of sediment. In con­ 22a ); propodus of fourth thoracic ap­ sequence, I have found it necessar y to assign pendage rectangular, generally longer the larval species on the basis of the adult than broad, and only slightly broader characteristics. 'If one has identified all adult than that _of fifth (Fig. 23/) C species occurring in a particular region, then, BB. Larvae hatch with biramous thoracic by studying the postlarval forms which show appendages (Fig. 25a) which undergo both adult and larval characteristics, it is po s­ transformation into typical subchelate sible to correlate the larval forms occurring form; propodus of fourth thoracic ap­ in the same region by working back from the pendage rounded, broader than long, adult species to ,the earliest recognizable lar­ more than twice as large as that of fifth vae of those species. Using this method, I (Fig . 27b) D have been able to identify larval stages of C. Telson with 4 or more intermediate den­ Pseudosquilla as those belonging to adult P. ticles between submedian and lateral ciliata. Larvae of Lysiosquilla and Squilla are denticles (Fig. 22b) . likewise assigned to their respective species, ·, Squilla (alima larva) but this identification is not positive. At this Cc. Telson with 3 or fewer intermediate den­ time larvae ofCoronida and Odontodactylus can­ ticles between submedian and lateral not be referred to any definite species . Larval denticles (Fig. 23e) : . forms of Gonodactylus have not been found in · . . .Pseudosquilla (pseuderichthus larva) these waters. D . Propodus of raptorial claw broad, dac­ The following key is designed to facilitate tylus slightly inflated at base (Fig. 26e) the identificatio n of the larvae with their re­ · . . . .Coronida (coroniderichthus larva) spective genera, and is intended to be used DD. Propodus of rapto rial claw lon g and for larvae in all stages of development. There slender, dactylus not inflated at base is no reference to species, but where the adult (Fig. 25b) . species is known it will be mentioned in the · . . . . .Lysiosquilla (lysioerichthus larva) description given for each genus. E. Spines of telson greatly elon gated (Fig. 28a) . Key to the Genera of Larval · . Odontodactylus (odonterichthus larva) Hawaiian Stomatopoda BE. Spines of telson not greatly elon gated · . . .. .Gonodactylus (gonerichthus larva) A. Larvae hatch without raptorial claw (an­ tizoea with biramous appendages on Squilla sp? , probably S. oratoria first 5 thoracic somites) or with rapto­ Fig. 22a- d rial claw (pseudozoea with only first and second thoracic appendages) and with STAGE 1 (Fig. 22a) : This is the earliest stage the propodus finely pectinate along its of the alima-type larva which has been found. upper margin; ischio-rneral articulation Larvae bear completely developed first and of raptorial claw always terminal in later second thoracic appendages, the second bein g stages (Figs. 22a, 25b ) B in the form of a raptorial claw; propodus of AA. Larvae always hatch with raptorial claw thi s claw with two small spines near its pro x­ (pseudozoea) and the propodus never imal end , upper margin finely pectinate; dac­ pectinate alon g its upper margin; ischio­ tylus nearly straight, no teeth along inner meral articulation of raptorial claw at a margin; eyes borne on long stalks, this con­ point in advance of the proximal end of dition bein g retained throughout successive the merus in later stages (Fig. 28a) . .E stages; antennules fully formed; antennal Hawaiian Stomatopods - TOWNSLEY 427

FIG. 22. Larval forms of Squilla. a, D orsal view of pseudo zoea, first abdominal appendage and telson (probably S. oratoria); b, do rsal aspecr of alima larva and firsr abdo minal appendage (probably S. oratoria); C, ventral aspect of alima larva (probably S. oratoria ); d, dor sal aspec t of alim a larva, first abdo minal appendage and urop od (Squilla sp.). scale present but flagella absent ; carapace very denticles and four between lateral and sub­ clear, covering nearly 0.5 the entire bod y, this median denticles. relation ship remaining the same throughout The specimens range in size from 5.5 to successive stages ; anterior margin of carapace 8.0 mm. from the tip of the rostral spine to trispinous, posterior margin with two pos­ the posterior margin of the telson . This stage terolateral spines and small zoeal spine be­ has been taken in plankton tows in the waters tween them ; lateral margins ofcarapace usual­ around Oahu throughout the year, most ly armed with three laterally directed and two abundantly during the summer months. ventrally directed denticles, small denticle at ST AGE 2 (Fig. 22b): This larval form is sim­ base of posterolateral spines; abdomen com­ ilar to the preceding stage except for a con­ pletely segmented , sixth abdominal somite spicuous change in size, the animals now indistinct; four pairs of biramous pleopods measuring 15 to 20 mm. Body proportions without gills, fifth pair represented by small remain the same, carapace still covering nearly buds; no uropods; telson longer than wide 0.5 total length of bod y; antennal flagella with 11 or 12 denricles between submedian present ; third , fourth , and fifth thoracic ap- 428 PACIFIC SCIENCE, Vol. VII, October, 1953 pendages present; sixth, seventh, and eighth Squilla sp? thoracic appendages represented by small Fig. 22d buds; abdominal appendages all present, with small gills attached to exopods; uropods be­ Approximately 200 specimens of this alima ginning to form; sixth abdominal somite with larval stage were taken in night light stations two small submedian spines; telson well at Honolulu Yacht Harbor, Oahu. They close­ developed, beginning to assume adult con ­ ly resemble the preceding larval types, but, as only one stage was collected , it is impossi­ dition with two well-developed spines on ble to assign the specimens to an adult species. distal margin and distinct submedian and in­ Carapace covers less than 0.3 total length of termediate denticles. bod y, very narrow; all appendages fully de­ As with the first stage, all the specimens veloped, gills represented by small buds on have been collected from regions where S. exopod of pleopods: telson very large com­ oratoria is either known or thought ro be pared with other bod y segments, composing present.Innumerable specimens have been about 0.25 total length of body; fully devel­ taken in plankton tows in the waters around oped uropods, bearing five spines along outer Oahu throughout the year. They may fre­ margin of exopod, with basal portion elon ­ quently be found in the stomach contents of gated into two spines , inner shorter than the yellowfin tuna, Neothunnus macropterus. outer. STAGE 3 (Fig. 22c): By the time the larvae Pseudosquilla ciliata have reached this stage, they have gained Figs. 23a-j, 24a-c further adult characteristics. Sixth, seventh, STAGE 1 (Fig. 23a): Brooks (1886) has il­ and eighth thoracic appendages fully devel-. lustrated a pseudozoea larva similar to this oped; uropods with adult characteristics , basal stage. This represents the earliest larval stage . portion elongated into two spines and small of members of the genus Pseudosquilla. By rounded lobe between them, exopods with observing the relative changes in the other eight spines , as yet immovable, along thei~ stages , I have come to the conclusion that outer margin; telson more adult-like in ap­ these are the youn g of P. ciliata. Typical pearance; no longitudinal carinae on body; erichthus larva with only first and second carapace still very transparent, no indication thoracic appendages present; abdomen with of facies of adult carapace showing through four pairs of biramous pleopods, without chitin. gills; telson nearly rectangular, with eight At this time the specimens measure 45 to small denticles between submedian denticles; 60 mm . in length. This stage forms a con­ eyes not borne on long st~lks as in alima of spicuous part of the plankton and has been Squil!« as articulation is close to ophthalmic taken in plankton tows from the surface down somite; antennules present ; antennae appear to a depth of 200 meters. as small buds ; merus and ischium of raptorial Th ere are no specimens beyond this stage claw with terminal articulation, propodus fine­ in the collections. It is rather unusual that ly pectinate along upper margin , dactylus no postlarval forms have been found in which composed of single spine; carapace trispinous the adult characteristics such as the bod y cari­ anteriorly, posteriorly bearing two long spines nae, the adult carapace, and the teeth on the reaching second abdominal somite; may Of raptorial dactylus are present. I assume from may not have small zoeal spine between two this that the larvae become bottom dwellers posterior spines ; carapace longer than broad , immediately after they lose the larval carapace. its lateral margins depressed . Hawaiian Stomatopods- TOWNSLEY 429

lJ f;JJ ~.•. ?'.... , c a

b

FIG. 23. Larval forms of Pseudosquilla ciliata (Fabricius). a , Dorsal aspect of pseudozoea and first abdominal appendage; b, lateral aspect of pseuderichthus larva, dorsal aspect of carapace and first abdominal appendage; c, dorsal aspect of last abdominal som ite, telson, and uropods of larva shown in b; d, e, lateral aspect of larger pseuderichthus larva and dorsal aspect of last abdominal somite, telson, and uropods; f, second, third, fourth, and fifth thoracic appendages of larva d.

Larval specimens in this stage of develop­ and nearly equal in size (not well shown in ment have been taken in great numbers in Fig. 23b); no appendages on sixth through plankton tows throughout the year at Kane­ eighth thoracic somites; eyes globular, pro­ ohe Bay, Oahu. Since adults of P. ciliata and truding from anterior margin of carapace; an­ oculata occur in abundance on the reefs in the tennae present; pleopods resemble those of bay, these specimens are believed to be the following stage, but have no gills; sixth ab­ first larval stage of a Pseudosquilla. Unfortu­ dominal somite distinct, bearing two sub­ nately, I find no way to differentiate the early median spines; uropods composed of short larvae of species in this genus . The specimens exopod and endopod, with basal portion elon­ vary from 3.5 to 5.0 mm. from the tip of the gated into two spines, inner shorter than rostral spine to the posterior margin of the outer; telson retains same general appearance telson . . of preceding stage, but somewhat larger. STAGE 2 (Fig. 23b, c): At this stage in de­ velopment the larvae show characteristics of Specimens measure 9.5 to 12.5 mm. in both the preceding stage and the fourth stage. length. Approximately 200 individuals in this Entire body and larval carapace have increased stage of development have been examined. in size but retain same relative proportions to Most ofthe specimens were taken in plankton one another; carapace mayor may not bear tows from Kaneohe Bay, Oahu, but they have small zoeal spine; first through fifth thoracic also been found in the tows from the Molokai appendages present; propodus of fourth and Channel made by the Territorial Division of fifth thoracic appendages rectangular in shape Fish and Game . 430 PACIFIC SCIENCE, Vol. VII, October, 1953

STAGE 3: This stage has not been found, STAGE 6 (Fig. 24a, b): This is the first but examination of the preceding stage and postlarval form of P. ciliata. No larval cara­ that of the next indicates what specimens in pace, but animals remain planktonic; eyes this stage of development should look like. resemble adult, nearlytubular in shape, cor­ By this time, all appendages should be pres­ nea followin g long axis of stalk; rostrum tri­ ent, gills should appear on the exopod of the angular ; carapace distinctly adult in shap e, pleopods, and the telson and uropods should with distinct gastric grooves, covering only be similar to the followin g stage. They should anterior 0.25 of body ; all appendages, except range from 15 to 20 mm. in length. second thoracic appendage, resemble those STAGE 4 (Fig. 23d-f ): Thoracic appendages ofadult; second thoracic appendage, raptorial all present, showing adult characteristics, ex­ claw, with shortened propodus and dact ylus cept raptorial claw; raptorial claw still very so that they fit into grooved merus and ischium long and slender, propodus much longer than 'when flexed, dactylus still lacks teeth on inner ischium and merus combined, dactylus com­ posed of a single long spine with no indica­ tion of teeth along inner margin; facies of postlarval carapace and rostrum show through chitin of larval carapace; urop ods greatly en­ larged with two immovable spines on outer margin of exopod, definite pits for others show through chitin, outer spine of basal prolongation of uiopods lon ger than innner; telson armed with two submedian, two inter ­ mediate, and two lateral spines, with 15 to 20 denticles between submedian and intermediate spine; facies of postlarval telson showing through chitin, with characteristic shape of adult, being nearly as broad as long. Specimens of this stage measure 20 to 22.5 mm . in length. Very few of these larvae have been collected in regions where the previous stages were most numerous. Occasionally the y are taken in the protected inshore waters, but they are more often found in the offshore waters such as the Molokai Channel. This seems to indicate that these larvae seek less sheltered waters and return to the shallow ~:. ·· · · ·· · · · { ' < ·· ~, . : ~ I...... •.:.. a inshore waters only when they lose the larval ~ carapace. fI5/ b STAGE 5: There are no great changes from the preceding stage at this time. Slight in­ crease in size; carapace very loosely attached to body; mandibular palp present, composed FIG. 24.Larval forms of Pseudosquilla ciliata (Fabri­ of three segments; nine or ten immovable cius). a ,b, D orsal aspect, second th oracic appendage, spines along outer margin of exopod of and first abdominal app endage of first posdarval stage; c, do rsal aspect of anterior segments of last posd arval uropod. . stage . Hawaiian Stomatopods- TOWNSLEY 431 margin; abdominal somites and appendages Brooks's specimens and those at hand agree distinctly adult in structure, gills on pleopods with those described by Claus (1872) from nearly fully developed, sixth abdominal so­ Messina. These very small larvae occur in mite with two well-developed submedian and great numbers in the ' plankton throughout two lateral spines; uropods with nine or ten the year. Due to their relatively small size movable spines along outer margin ofexopod, (2.5 to 3.5 mm. ) they are very often over­ spines of elongated basal portion changing looked or cast aside as some unknown form. relationship to one another so that in some Although they lack the characteristic sub ­ specimens these two spines are nearly equal chelate appendages, the eyes and telson have in length; telson with three distinct longi­ distinct stomatopod characteristics. These lar­ tudinal carinae, median and two submedians, vae represent the first larval stage of Lysio­ posterior margin with pair of movable sub­ squilla or Co ronida, and because of their great median spines bearing series of denticles be­ abundance I am inclined to believe that they tween them , pair of immovable intermediate belong to Lysiosquilla which is apparently spines with two denticles between them and more abundant in Hawaiian waters than Coro­ submedian spines, pair of immovable lateral nida. There is no diagnostic character suffi­ spines. ciently developed to indicate conclusively the The specimens now reach a maximum species to which these specimens belong, and length of 20 to 25 mm. This developmental none have ever been reared in captivity to the stage is very common in the plankton during following stage. The carapace completely the spring and summer months and occurs in covers the animal from head to abdomen both the inshore and offshore waters of Ha­ leaving only the eyes and telson exposed. waii. They may occasionally be found in the There are five pairs of biramous thoracic ap­ stomach contents of the yellowfin tuna, N eo­ pendages present , the abdomen is only faintly thunnus macropterus, and the skipjack, Katsu­ segmented, and the pleopods are absent. This wonuspelamis. larval form gives rise to an erichthus in later STAGE 7 (Fig. 24c): There is little change stages. in this form from the preceding stage. Ros­ trum nearly adult in form, broader than long; Lysiosquilla sp? eyes more tubular in shape; raptorial dactylus Figs. 25b, c composed ofsingle tooth, facies ofother teeth on inner margin can be seen through chitin Nine specimens of this lysioerichthus stage in some specimens. were collected in a plankton tow in the Molo­ Individuals in this developmental stage kai Channel. .In all respects this larval form have been taken in the plankton from Ka­ represents the typicallysioerichthus. No spines neohe Bay, Oahu, and at Honolulu Yacht between submedian and lateral spines on tel­ Harbor, Oahu. This indicates that the larvae son (Fig. 25c); raptorial propodus finely pec­ are returning to the shallower, more pro­ tinate along upper margin , dactylus composed tected waters and are preparing to settle of single acute spine, not inflated at base, down on the bottom to complete their life ischio-meral articulation terminal; all thoracic cycle. appendages present , propodus of third and Lysiosquilla sp? fourth rounded and broader than long, fourth nearly twice as large as that of fifth; abdomen Fig.25a increasing in breadth posteriorly; pleopods Brooks (1886) has illustrated this antizoeal all present, gills absent; uropods partially de­ form and states that numerous specimens were veloped, exopod and endopod both present, taken by the "Challenger" at Honolulu. Both with basal portion elongated into two spines, 432 PACIFIC SCIENCE, Vol. VII, October, 1953

b c

d

FIG. 25. Larval forms of Lysiosquill«. a, Aneizoea; b, lateral aspect of lysioerichthus larva and first abdominal appendage; c, dorsal aspect of last abdominal somite, telson, and uropods of same ; d, vent ral aspect of lysioerich­ thus larva; e, ventral aspect of last larval stage ofL. maculata; f, dorsal aspect of first pos clarval stage of L. maculata. inner longer than outer; carapace nearly 'as opening of carapace is completely closed . broad as long from base ofanterolateral spines Specimens measure 14.5 to 28.5 mm . in to base of posterolateral spines, lateral mar­ length . The 28.5 mm. specimen shows the gins turned downward and toward median facies of ten spines along the inner margin of line; most preserved specimens with abdomen the raptorial dactylus much the same as those and telson reflexed anteriorly so that ventral seen in the adults of L. macu!ata. Hawaiian Srornatopods- TOWNSLEY 433

Lysiosquilla sp ? Fig. 25d This lysioerichthus larval form is represent­ ed by a single specimen taken in a plankton tow in the Molokai Channel. Its very large a size (45 mm.) immediately draws attention. All structures are the same as for the previ­ ously described larvae of this genus. It was temporarily believed to belong to L. maculata, but its size and relative immaturity (d . Fig. 25e, which is onl y 15 mm . long) seem to indicate that it may be the larval stage of a species of Lysiosquilla which has not been fourid in the adult form in Hawaii. c

Lysiosquilla maculata Fig. 25e,I Figure 25e represents one of seven lysio­ erichthus larvae, three ofwhich were collected in plankto n tows at Tru k Island and four from Kawaihae, Hawaii. These are fully de­ veloped larvae, which I believe t o be the last larval stage of L. maculata. This conclusion is based on an examinat ion of postlarval forms d and the adults of this species. Appendages, except raptorial claw, telson, and uropods, FIG. 26. Larval forms of Coronida sp. a, Lareral aspect resemble those of adult L. maculata; facies of last larval stage of coroniderichthus larva; b, dorsal aspect of last abdominal somite, telson, and uropod s of postlarval telson show through chitin , very of same ; c, first abdominal appendage of same; d, do r­ closely resembling telson of postlarval forms sal aspect of anterior segments of first posdarval stage; shown in Figure 25f It is very probable from e, second thoracic appendage of same. these facts that the next molt would prod uce the postlarval stage of L. maculata. Specimens pods. A single specimen measurin g 16.5 mm. range from 15 to 16 mm. from the tip of the in length was collected at Kawaihae, Hawaii, rostral spine to the posterior margin of the at a night light station, which further sub­ telson. stantiates the identification because the adult The postlarval form (Fig. 25/ ) undoubtedly species is very common in that region. belongs to L. maculata. All anterior append­ ages are the same as those of the adult, and Coronida sp? the raptorial dactylus bears nine teeth on its Fig. 26a-e inner margin similar to the con dition found in some of the adults of this species. The In Figure 26a there is represented the last telson .and urop ods have the same general larval stage (coroniderichthus) of some un­ appearance of the adult as well. These spec­ known species of Coronida. This form is imens and the adults have eight spines along found in the Ala Wai Canal, Oahu, and at the outer margin of the exopod of the uro- Kawaihae, Hawaii, in moderate numbers dur- 434 PACIFIC SCIENCE, Vol. VII, Ocrober, 1953

equal numbers with the previous stage. Only the carapace and anterior segments and the raptorial claw have been illustrated as all other structures are the same as those seen in the preceding stage. The inner margin of the dac­ tylus is armed with ten teeth. There' is no distinct increase in size over that of the last larval stage. These forms may be larval stages of C. sinuosa, but the larval structures bear no resemblance to diagnostic characteristics ofthis species. Apparently the species to which these larvae belong is quite common, al­ though it has not been found as yet.

Coronida sp? Fig. 27a- c Five coroniderichthus larvaemeasuring 18 I mm to 20 mm. from the tip of the rostral spine to the posterior margin of the telson were collected at Kawaihae, Hawaii. The append­ ages indicate that they belong to some species of Co ronida, but no adult specimens have been foun d which have the same characteristics as c those possessed by this larval stage. I believe , that the first 'postlarval stage would appear FIG. 27. Larval form of Coronida sp. a, D orsal aspect at the next molt because the facies of the of coroniderichthus larva ; b, second, first, third, fourth, and fifth thoracic appendages; c, first abdominal ap­ adult carapace, telson, and raptorial dactylus pendage. show through the very clear chitin. These specimens are unique in that the larval cara­ ing the summer. Larvae resemble lysioerich­ pace covers all but the l ast two abdominal thus in shape of tho racic appendages, but somites and the telson, and the median an­ with raptorial dactylus slightly inflated at its terior spine and posterolateral spines are base; propodus of fourth tho racic appendage greatly elongated . This condition is not foun d rounded, nearly twice as large as that of fifth in any other larval form, hence it should which is rectangular; all abdominal append­ serve as a diagnostic character for the separa­ ages present; no gills; telson rectangular as tion of this larval form from others belonging in most larval forms , bearing two denticles to this genus. between submedian and lateral denticles ; uro­ pods present, outer margin of exopod armed Odontodactylus sp? with three spines, basal portion elongated Fig. 28a-e into two spines, outer longer than inner . Specimens measure 10.0 to 14.5 mm . from Eight odonterichthus larvae, rangin g in the tip of the rostral spine to the posterior length from 18 to 35 mm., were taken in a margin of the telson . plankton tow in the Molokai Channel. They Figures 26d, e represent.the first postlarval resemble pseuderichthus larvae very much, stage of this species which is found in nearly and it is possible to distin guish the two only Hawaiian Stornatopods- TOWNSLEY 435

FIG. 28. Larval forms of Odontodactylus. a , Lateral aspect of odonrerichthus larva, first abdominal appendage, and dorsal aspect of last abdo minal somite, telson, and uropods; b, lateral aspect of odonterichthus larva and dorsal aspect of last abdo mi nal somite, telson, and uropods ; c, first, third, fourth, and fifth thoracic appendages of same ; d, second thoracic appendage of same ; e, first abdominal appendage of same. . by comparing the raptorial claws. In pseu­ parisons with their figures show that the Mo­ derichthus larvae this appendage is finely lokai Channel specimens definitely belon g to pectinate along the upper margin of the pro­ the odonrerichthus type. Foxon (1932) uses podus, and the dactylus is not inflated at the the character of the long spines found on the base. In odonrerichthus, however , the rapto­ telson of Odontodactylus larvae for separating rial propodus is not pectinate along its upper them from Gonodacytlus, although he does not margin, and the dactylus is inflated at its present any figures to illustrate his statements. base. In later stages the ischio-meral articula­ Since no Gonodactylus larvae have been col­ tion of the raptorial appendage would be a lected in the Hawaiian waters, I cannot say useful diagnostic character, but none of the whether or not this is a valid character for the specimens has attained this condition, which separation of the larvae of the two genera, but is present in the adult. This genus is un­ comparisons with Claus's and Brooks 's figures doubtedly closely related to Gonodactylus, but of gonerichthus larvae show that the spines it has been impossible to compare the larvae on the telson of their specimens are consider­ of the two genera because no specimens of ably shorter than those found in the present the latter have been collected in Hawaiian specimens. With this in mind I have placed waters. Claus (1872) and Brooks (1886) have the character in the key with the hope that both illustrated gonerichthus larvae which -sorne future worker who may find the go­ were collected from other regions, and com- nerichthus larvae will be saved some effort in 436 PACIFIC SCIENCE, Vol. VII, October, 1953

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