(Crustacea: Cryptochiridae)! Occurring in the Pacific Ocean

Revision of the Genera of Gall Crabs (Crustacea: Cryptochiridae)! Occurring in the Pacific Ocean

Roy K. KROPP2

ABSTRACT: The coral gall crabs, Family Cryptochiridae, occurring in the Pacific Ocean are reviewed. Fifteen genera, including four new genera, are recognized: Cryptochirus Heller; Dacryomaia, new genus; Fizesereneia Takeda & Tamura; Fungicola Serene; Hapalocarcinus Stimpson; Hiroia Takeda & Tamura; Lithoscaptus Milne Edwards; Neotroglocarcinus Takeda & Tamura; Opecarcinus Kropp & Manning; Pelycomaia, new genus; Pseudocryptochirus Hiro; Pseudohapalocarcinus Fize & Serene; Sphenomaia, new genus; Utinomiella Kropp & Takeda; and Xynomaia, new genus. Host, depth, and distribution records are given for each genus. A key to the females of all gall crab genera is included.

THE CORAL GALL CRABS, Family Crypto ramous or biramous) of the second pleopod chiridae, are obligate associates of living of the female and the host coral taxon on scleractinian corals. Not only do they reside which the crab species occurred. McCain and in galls, tunnels, or pits in the coral skeleton, Coles (1979) found the second pleopod to be but they also feed on the host coral mucus and uniramous on one side of the abdomen of an tissues (Kropp 1986). Though the family has individual, biramous on the other, and vari been known for 130 yr, its taxonomy has been able among individuals of Utinomiella dimor marred by many errors. These errors have pha (Henderson 1906). I have made similar resulted from failures to consider type mate observations for this and other species among rial or available literature, use of inappro my collections from Micronesia. Crypto priate characters to define taxa, or failure to chirids show some degree of host specificity, follow proper taxonomic procedures. which some authors thought could be used to Two problems related to failure to consider define genera (Fize and Serene 1957, McCain type material were resolved by the reestab and Coles 1979). Both features subsequently lishment of Lithoscaptus Milne Edwards and were shown to be unreliable (Kropp and Cryptochirus hongkongensis Shen, the latter as Manning 1987). the type species of Neotroglocarcinus Takeda In a monograph ofthe gall crabs from Viet & Tamura (Kropp 1988a,b). nam, Fize and Serene (1957) created a new Two characters inappropriate for use in the genus, Neotroglocarcinus, and initiated the definition of genera were the nature (uni- use of subgenera in the genus Troglocarcinus Verrill. The subgenera they established were based on the family of corals on which the 1 Contribution no. 284 from the University of Guam crabs were found. However, they erred in Marine Laboratory. Financialsupport was provided by creating the subgenus Mussicola by including theUniversity of Maryland Department ofZoology and within it T. corallicola Verrill, the type species Graduate School, a grant to G. J. Vermeij by the Bio logical Oceanography Section of the National Science of Troglocarcinus. They further erred by not Foundation, and the Lerner-Gray Fund for Marine naming type species for any of their newly Research. This is from a dissertation submitted to the created genus-group taxa. Thus, the new Graduate School, University of Maryland, in partial ful genus-group names were unavailable. fillment of the requirements for thePh.D.degree in zoo logy. Manuscript accepted 10 February 1990. Serene (1966) made two generic names 2 Presentaddress: Battelle Ocean Sciences, 1431 Spin available when he designated Cryptochirus naker Drive, Ventura, California 93001. rugosus Edmondson as the type species of 417 418 PACIFIC SCIENCE, Volume 44, October 1990

Favicola and Troglocarcinus utinomi Fize & and 1986. Specimens from these collections Serene as the type of Fungicola. His actions have been deposited in the National Museum also helped establish the host taxon as a gene of Natural History, Smithsonian Institution ric character. (NMNH), Washington, D.C.; the Bernice P. By the beginning of the 1980s the taxono Bishop Museum (BPBM), Honolulu, Hawai'i; mic situation of the family was far from and the Natural History Museum of Los serene, but the work of Takeda and Tamura Angeles (LACM), Los Angeles, California. in Japan had begun to unravel some of the Supplementary material was borrowed from confusion. Takeda and Tamura (l981b) re the BPBM; the NMNH; the British Museum moved Pseudocryptochirus as a "catch all" of Natural History (BMNH), London, Eng taxon by creating two new genera, Hiroia and land ; the Museum National d'Histoire Natu Utinomia, for some species previously as relle (MNHN), Paris , France; the Naturhis signed to Pseudocryptochirus. Utinomia torisches Museum (NMW), Vienna , Austria; Takeda & Tamura, 1981 was later determined and the National Science Museum (NSMT), to be a junior homonym of Utinomia Tom Tokyo, Japan. I examined the types of many linson , 1963, a genus of acrothoracican bar species; these are marked by an asterisk in the nacle, and was replaced by Utinomiella Kropp lists of species. & Takeda 1988. Most of the figures are based on material Takeda and Tamura (1980b) also erected a from Micronesia and were prepared with the new genus, Fizesereneia, to accommodate use of a camera lucida on a Wild M-5 dissec species placed in the erroneously created Mus tion microscope or were traced from scanning sicola Fize & Serene, 1957. However, they did electron microscope (SEM) micrographs. The not realize that Mussicola could be used as type species of each genus is figured. Setae on an available name by designating one of the the surfaces of the carapace and pereopods species in it as its type. Kropp and Manning are not shown. Specimens for SEM study were (1987) rectified this by naming T. corallicola prepared as described in Kropp (1986). as the type species of Mussicola, rendering I have restricted the synonymies to the first Mussicola a junior synonym of Troglocar usage ofa particular name for a taxon, includ cinus. Takeda and Tamura (1980b) further ing erroneous spellings. Abbreviations used in erred by misidentifying the type species of the text or figures are as follows: m, meters; their new genus Fizesereneia. Kropp (l988c) mm, millimeters; MXL, maxilla; MXP, maxil petitioned the International Commission on liped; P, pereopod; PLP , pleopod; and TL, Zoological Nomenclature to confirm Troglo type locality. carcinus heimi Fize & Serene, 1956 as the type In the diagnoses the male P-1 is described species of Fizesereneia . as robust if the height of the manus is sub However, many morphological inconsis equal to the length of its dorsal margin, slen tencies within the generic structure of the der if the height ofthe manus is much less than family still remained by the mid-1980s. Here, the length of its dorsal margin. Similarly, the I use morphology to reorganize the generic propodus ofleg P-2 of the female is described placement of the species of gall crabs occur as thick if its height is subequal to its length; ring in the Pacific, removing totally the con slender if the length is 1.5 or more times the cept of host affinity as a generic character. I height. The median indentation of the an also pro vide a key to the females ofall known terior margin of the epistome is characterized cryptochirid genera . The species of gall crabs as narrow if its maximum width is less than occurring in the Atlantic were reviewed by half or wide if its maximum width is greater Kropp and Manning (1987). than half the width of the anterior margin. Also, the median indentation is termed shal low if its depth is less than half or deep if its depth is greater than half the length of the MATERIALS AND METHODS epistome. In some genera the median inden I collected much ofthe material used in this tation is barely noticeable and is termed revision in Micronesia and Hawai'i in 1984 slight. Revision of Pacific Gall Crab Genera-i-Knorr 419

I have listed only those host, depth, and dicates unpublished data from my collections distribution records that I am reasonably cer in Micronesia and Hawai'i. tain are accurate. The acronym "RKK" in-

KEY TO GENERA OF GALL CRABS , FAMILY CRYPTOCHI RIDAE (Based on Females) I. Carapace deflected anteriorly ...... 2 I. Carapace not deflected anteriorly...... 10 2. Anterior extension of sternite of P-I smooth...... 3 2. Anterior extension of sternite of P-I with granules or tubercles ...... 6 3. Anterior carapace with two bowl-shaped concavities 4 3. Anterior carapace without bowl-shaped concavities...... 5 4. Distal margin of antennal segment 2 with lateral spine; pterygostomial region not fused to carapace...... Pelycomaia Kropp, new genus 4.Distal margin of antennal segment 2 without lateral spine; pterygostomial region fused to carapace Pelycomaia 5. Pterygostomial region fused to carapace; P-2 merus lacking distomesial expansion . . ... · Lithoscaptus 5. Pterygostomial region not fused to carapace; P-2 merus with distomesial expansion . . .. · Troglocarcinus (Atlantic) 6. P-2 merus with distomesial expansion 7 6. P-2 merus without distomesial expansion ...... 8 7. Pterygostomial region fused to carapace; P-4 sternite with median suture . ·...... Opecarcinus 7. Pterygostomial region not fused to carapace; P-4 sternite without median suture . ·...... Neotroglocarcinus 8. Midportion of carapace with clusters ofrounded tubercles...... Cryptochirus 8. Midportion of carapace with isolated tubercles and granules ...... 9 9. Carapace widest at midlength; anterior carapace without deep inverted V-shaped groove · Sphenomaia Kropp, new genus 9. Carapace widest anterior to midlength; anterior carapace with deep V-shaped groove · Dacryomaia Kropp, new genus 10. Pterygostomial region fused to carapace ...... II 10. Pterygostomial region not fused to carapace...... 16 11. P-3 coxa with well-developed anterior lobe...... 12 11. P-3 coxa anterior lobe lacking or reduced ...... 13 12. MXP-3 with exopod Fungicola 12. MXP-3 lacking exopod ...... Utinomiella 13. Carapace about as long as wide ...... 14 13. Carapace longer than wide ...... 17 14. P-4, P-5 coxal lobes well-developed Pseudohapalocarcinus 14. P-4, P-5 coxal lobes absent Hapalocarcinus IS. MXP-3 with exopod Hiroia 15. MXP-3 lacking exopod Detocarcinus (Atlantic) 16. Merus of P-2 with distomesial expansion; distal margin of antennal segment 2 lacking lateral spine...... 17 16. Merus of P-2 without distomesial expansion; distal margin of antennal segment 2 with lateral spine Xynomaia Kropp, new genus 17. Anterior extension ofP-1 sternite with granules Pseudocryptochirus 17. Anterior extension of P-I sternite smooth Cecidocarcinus (Atlantic) 420 PACIFIC SCIENCE, Volume 44, October 1990

SYSTEMATIC ACCOUNT hood. P-I cutting edges entire. P-2 merus lack ing distomesial projection; propodus slender. Family CRYFTQCHIRIDAE Paulson, 1875 P-3 coxa with reduced, P-4 coxa with well developed anterior lobe. P-5 dactylus rotated Lithoscaptes Milne Edwards, 1862:FIO [ver anteriorly. PLP-3 of female uniramous. Male nacular name; type genus Lithoscaptus with robust P-l; abdomen length 3.3 times Milne Edwards, 1862]. width, segments subequal in width, lateral Cryptochirinae Paulson, 1875: 72 [po 78 in margins parallel. PLP-l curved laterally, translation; type genus Cryptochirus Heller, apex pointed, lateral margin with simple 1861]. setae, mesial margin with few pappose setae Cryptochiridae.- Richters, 1880: 159;Kropp proximally. and Manning, 1985:954. Lithoscaptidae.-Richters, 1880: 159. REMARKS: Fize and Serene (1957) failed to Hapalocarcinidae Calman, 1900:49 [type designate a type species for Troglocarcinus genus Hapalocarcinus Stimpson, 1859]. (Favicola), thus rendering Favicola Fize & Hapalogarcinidae.c-Coelho & Ramos, Serene unavailable for use as a genus-group 1972: 205 [erroneous spelling]. name [International Code of Zoological No menclature (lCZN) Art. 13(b)]. Serene (1966) Genus Cryptochirus Heller, 1861 designated Cryptochirus rugosus Edmondson as the type species ofFavicola. Kropp (I988b) Cryptochirus Heller, 1861: 19 [type species: determined Cryptochirus rugosus to be a Cryptochirus coralliody tes Heller, 1861: 19, junior synonym of C. coralliodytes. by monotypy; gender masculine]. Cryptochyrus.-Nobili, 1906: 325 [erroneous HOSTS: Faviidae-Cyphastrea, Barabat- spelling]. toia, Favia, Favites , Goniastrea, Leptoria, Favicola Fize & Serene, 1957: 84 [name un Montastrea, Platygyra (see Fize and Serene available]. 1957, RKK). Oculinidae-Cyathelia (see Favicola Serene, 1966:396 [type species: Takeda and Tamura 1983). Cryptochirus rugosus Edmondson, 1933: 6, DEPTH: < 1 to 30 m (Takeda and Tamura by original designation; gender masculine]. 1983, RKK). DIAGNOSIS: Carapace rectangular, longer than broad, widest near midlength, convex in DISTRIBUTION: Red Sea (Heller 1861);Viet lateral view, deflected anteriorly; mesogastric nam (Fize and Serene 1957); Japan (Takeda region inflated. Posterior region of carapace and Tamura 1983); Micronesia-Palau, isolated by system ofgrooves, cardiointestinal Guam, Pohnpei (RKK). region outlined. Pterygostomial region fused to carapace. Epistome with parallel lateral Cryptochirus coralliodytes Heller, 1861 ridges, anterior margin sinuous with shallow, Figure 1 wide median indentation. Lateral lobe of antennule elliptical, extending beyond eye Cryptochirus coralliodytes Heller, 1861: 19 stalk. Antennal segment 2 longer than broad, [TL: Red Sea; lectotype* NMW] . distal margin lacking lateral spine. MXP-3 Cryptochirus rugosus Edmondson, 1933:6 with exopod; merus with distolateral projec [TL: Washington Island (now known tion, mesial margin with setae. Inner surface as Teraina, Kiribati); holotype* BPBM of second segment of MXP-2 endopod with S3668]. setae at distal margin. MXP-l endopod trian Troglocarcinus (Favicola) rugosus.-Fize & gular, widest distal to midlength; distal mar Serene, 1957: 85. gin convex. Ventral thorax longer than wide, Favicola rugosus.-Serene, 1966: 396. concave mesially. Anterior extension of ster Favicola rugosum.-Lund0er, 1974: 10. nite of P-l with granules. Sternite of P-4 with Favicola rugosa.- Takeda & Tamura, 1981 a : suture. Female gonopore oval, with anterior 43. h

FIGURE I. Cryptochirus coralliodytes Heller. Female lectotype (Red Sea): a, dorsal view of carapace; b, lateral view; c, thoracic sternites ; d, P-2; e, epistome;j, MXP-3; g, endopod of MXP-l; h, MXL-l. Male (Guam): i, abdomen; j , PLP . Scale: b = 2.1 mm; c = 1.0mm;a = 0.8 mm;d, i = 0.7 mm ;f= 0.4 mm; e = 0.3 mm;g,j = 0.2mm;h = 0.1 mm. 422 PACIFIC SCIENCE, Volume 44, October 1990

Cryptochirus planus (Takeda & Tamura, REMARKS: Dacryomaia is most closely re 1983), new combination lated to Hiroia (see Kropp 1988d), but is dis Favicola plana Takeda & Tamura, 1983: 4 tinguishable from it by having the anterior [TL Koza, Japan; holotype* NSMT-Cr portion of the carapace deflected and by hav 8556]. ing a well-developed anterior lobe on the coxa of P-3. In Hiroia the anterior carapace is not deflected and the coxa ofP-3lacks an anterior Genus Dacryomaia Kropp, new genus lobe. Two early works by Fize and Serene Dacryomaia Kropp, new genus [type species: (1956a ,b) have been frequently cited as Fize Cryptochirus edmonsoni Fize & Serene, and Serene (1955a,b) . Fize and Serene (1957) 1956b: 379, by original designation; gender used both dates: 1955 in the text (p. 48 and feminine]. others) and 1956 in the bibliography (p.183). However, Ruth Cooper (pers. comm., 1988), a DIAGNOSIS: Carapace rectangular, longer than broad, widest anterior to midlength, con zoologist with the International Commission on Zoological Nomenclature, informed me vex in lateral view, deflected anteriorly; with that the actual publication date ofParts 5 and deep inverted V-shaped groove flanking me 6 was 31 March 1956. Therefore, all taxa des sogastic region, latter inflated. Cardiointes cribed in those articles should be dated 1956. tinal region outlined by depression. Pterygo stomial region fused to carapace. Epistome HOSTS: Thamnasteriidae-Psammocora (see with parallel lateral ridges, anterior margin Fize and Serene 1957). Siderastreidae straight with slight median indentation. Lat Coscinaraea (RKK). Faviidae-Cyphastrea, eral lobe of antennule oval, extending to or Goniastrea, Leptastrea (see Takeda and Ta just beyond eyestalk. Antennal segment 2 mura 1981a, RKK). Fize and Serene (1957) longer than broad, distal margin lacking listed Pavona (Agariciidae) as a host for D . spine. MXP-3 with exopod; merus with disto edmonsoni. However, the host was figured and lateral projection, mesial margin with setae. the crescentic pit made by the crab is typical Inner surface of second segment of MXP-2 for species of Opecarcinus, not D. edmonsoni. endopod with setae at distal margin. MXP-l Therefore, I think that host record was an endopod triangular, widest at midlength; error. distal margin evenly convex. Ventral thorax longer than broad, flat. Anterior extension of DEPTH: < 1 to 8 m (RKK). sternite of P-I with transverse band of gran DISTRIBUTION: Vietnam (Fize and Serene ules. Sternite ofP-4 with suture. Female gono 1957); Japan-lzu Islands, Ogasawara Is pore elliptical, with anterior hood. P-I dac lands, Ryukyu Island s (Takeda and Tamura tylus with low tooth proximally on cutting 1980d, 1981a, 1983); Micronesia-Palau, edge. P-2 merus lacking distomesial projec Guam (RKK). tion; propodus thick. P-3, P-4 coxae with well developed anterior lobes. P-5 dactylus rotated Dacryomaia edmonsoni (Fize & Serene, 1956), anteriorly. PLP-3 offemale uniramous. Male new combination with robust P-l; abdomen length 2.6 times Figure 2 width , segment 3 widest, lateral margins sub parallel. PLP-l slightly curved laterally, apex Cryptochirus edmonsoni Fize & Serene, 1956b: pointed, lateral margin with several simple 379 [TL: Nhatrang, Vietnam; location of setae, with few pappose setae proximally, type unknown]. mesial margin with few simple setae proxi mally. Dacryomaia japonica (Takeda & Tamura, 1981), new combination ETYMOLOGY: From the Greek, dakryon , meaning droplet, in combination with maia, a Favicolajaponica Takeda & Tamura, 1981a: kind of crab, in reference to the shape of the 47 [TL: Kuroshima, Ryukyu, Japan; holo tubercles on the anterior part of the carapace. type* NSMT-Cr 7422]. FIGURE 2. Dacryomaia edmonsoni (Fize & Serene). Female (Guam: a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;j, MXP-3; g, endopod of MXP -I ; h, MXL-1. Male (Guam): i, abdomen;j, PLP . Scale: a- c, i = 1.0 mm; d = 0.5 mm;!= 0.3 mm; e = 0.2 mm; g, h,j = 0.1 mm. 424 PACIFIC SCIENCE, Volume 44, October 1990

Genus Fizesereneia Takeda & Tamura, 1980 ing a hexagonally shaped carapace, and by being a different color. The type of T. heimi Fizesereneia Takeda & Tamura, 1980b: 137 (see Fize and Serene 1957: fig. 29A) has the [type species: Troglocarcinus heimi Fize & median ridge completely dividing the anterior Serene, 1956a: 378, pending confirmation concavities and has a rectangular carapace. by the International Commission on Zoo The color of T. heimi that I collected in Guam logical Nomenclature (Kropp 1988c); gen was predominately gray, matching that fig der feminine]. ured by Fize and Serene (1957: pI. 18B) and Fizeserenia.- Kropp & Manning, 1987: 2 differing from the predominately green color [erroneous spelling]. of the undescribed species, also figured by DIAGNOSIS: Carapace hexagonal, longer Fize and Serene (1957: pI. 18A). Because the than broad, widest anterior to midlength, con genus was based on a misidentified type spe vex in lateral view, deflected anteriorly, cies, I petitioned the International Commis formed into 2 bowl-shaped concavities; meso sion on Zoological Nomenclature to confirm gastric region inflated. Cardiointestinal re T. heimi Fize & Serene as the type species of gion flanked by 2 depressions. Pterygostomial Fizesereneia (see Kropp 1988c). region not fused to carapace. Epistome with parallel lateral ridges, anterior margin sin HOSTS: Mussidae-Lobophyllia , Symphyl uous with shallow, wide median indentation. lia, Acanthastrea (see Fize and Serene 1957, Lateral lobe of antennule oval, extending be Takeda and Tamura 1980b). yond eyestalk. Antennal segment 2 longer DEPTH: 1 to 15 m (Takeda and Tamura than broad, distal margin with lateral spine. 1980b, RKK). MXP-3 with exopod; merus with distolateral projection, mesial margin with setae. Inner DISTRIBUTION: Vietnam (Fize and Serene surface of second segment of MXP-2 endo 1957);Indonesia (Serene et al. 1974);Japan pod with setae at distal margin. MXP-l en Izu Islands, Ryukyu Islands (Takeda and dopod triangular, widest distal tomidlength; Tamura 1980b); Australia (McNeill 1968); distal margin convex. Ventral thorax longer Micronesia-Palau, Guam, Pohnpei (RKK). than wide, flat. Anterior extension of sternite of P-l smooth. Sternite of P-4 with suture. Fizesereneia heimi (Fize & Serene, 1956) Female gonopore elliptical, with anterior Figure 3 hood. P-I cutting edge entire. P-2 merus lack Troglo carcinus heimi Fize & Serene, 1956a : ing distomesial projection; propodus slender. 378 [TL: Nhatrang, Vietnam; location of P-3, P-4 coxae with moderate anterior lobes. type unknown]. P-5 dactylus rotated anteriorly. PLP-3 of fe Troglocarcinus (Mussicola) heimi. -Fize & male uniramous. Male with robust P-l; abdo Serene, 1957: Ill. men length 2.3 times width , segment 4 widest, Fizesereneia heimi. - Takeda & Tamura, lateral margins convex. PLP-l curved later 1980b: 137. ally, apex pointed, lateral margin with many simple setae, mesial margin with few simple Fizesereneia ishikawaiTakeda & Tamura, 1980 setae proximally. Fizesereneia ishikawai Takeda & Tamura, REMARKS: Takeda and Tamura (I980b) 1980b: 144 [TL: Ishigakijima Island , designated T. heimi as the type species of Ryukyu Islands, Japan; holotype* NSMT Fizesereneia, a genus named in honor of the Cr 6340]. two Gaul crab specialists. However, the spe cies they described and figured was not T. Fizesereneia stimpsoni (Fize & Serene, 1956) heimi, but an undescribed species. Their fig ured species differs from T. heimi by having Troglocarcinus stimpsoni Fize & Serene, the median ridge incompletely separating the 1956b: 380 [TL: Nhatrang, Vietnam; loca concavities on the anterior carapace, by hav- tion of type unknown]. FIGURE3. Fizesereneia heimi (Fize & Serene). Female (Guam): a, dorsal viewof carapace; b,lateral view; c, thoracic sternites; d, P-2; e, epistome;j, MXP-3; g, endopod ofMXP-I; h, MXL-I . Male (Guam): i, abdomen;j, PLP. Scale: a, b, d, i = 1.0 mm; c = 0.9 mm; e = 0.3 mm; g, h, j = 0.1 mm. 426 PACIFIC SCIENCE, Volume 44, October 1990

Troglo carcinus (Mussicola) stimpsoni.-Fize available [ICZN Art. 13(b)] until Serene & Serene, 1957: 116. (1966) designated T. utinomi as the type spe Fizesereneia stimpsoni.- Takeda & Tamura, cies for the genus. 1980b: 146. Serene (1966), and later Takeda and Ta mura (1979), emended the specificname utino mi to utinomii. These actions were improper, Genus Fungicola Serene, 1966 as the ICZN [Art. 31(a) (ii)] specifies that the Fungicola Fize & Serene, 1957: 122[name un stem ofa species-group name is determined by available]. the action of the original author. Fize and Fungicola Serene, 1966: 396 [type species: Serene (1956a) originally used utinomi, thus Troglo carcinus utinomi Fize & Serene, defining the stem as "utinom," 1956a :377, by original designation; gender I examined the holotype, a male, ofPseudo masculine]. cryptochirus ishigakiensis and determined that Fungicora.-Takeda & Tamura, 1986:64 it differed from some of the males of F. uti [erroneous spelling]. nomi only by having a less convex abdominal outline. However, this condition occurred in DIAGNOSIS: Carapace rectangular, longer some other males of F. utinomi that I ex than broad, widest anterior to midlength, flat amined, thereby making it of dubious value in lateral view, not deflected anteriorly, with for distinguishing the two species, much less broad inverted U-shaped depression; meso placing them in separate genera. gastric region slightly inflated. Cardiointes tinal region outlined. Pterygostomial region HOSTS: Fungiidae-Fungia, Podobacia, fused to carapace. Epistome with parallellat Sandalolitha (as Parahalomitra) (see Fize and eral ridges, anterior margin sinuous with shal Serene 1957, Monod and Serene 1976). low, wide median indentation. Lateral lobe DEPTH: 1 to 15 m (RKK). ofantennule oval, extending just beyond eye stalk. Antennal segment 2 longer than broad, DISTRIB UTION : Vietnam (Fize and Serene distal margin lacking lateral spine. MXP-3 1957); Indonesia (Serene et al. 1974); Japan with exopod, merus with distolateral projec Ryuk yu Islands (Takeda and Tamura 1979); tion, mesial margin with setae. Inner surface Micronesia-Palau, Guam (RKK). of second segment of MXP-2 endopod with setae at distal margin. MXP-l endopod trian Fungicola utinomi (Fize & Serene, 1956) gular, widest distal to midlength, distal mar Figure 4 gin convex. Ventral thorax longer than broad, Troglocarcinus utinomi Fize & Serene, 1956a: flat. Anterior extension ofsternite of P-I with 377 [TL: Nhatrang, Vietnam ; location of few granules. Sternite of P-4 with suture. Fe type unknown] . male gonopore oval, with anterior hood. P-l Troglocarcinus (Fungicola) utinomi.-Fize & cutting edges entire. P-2 merus without dis Serene, 1957: 124. tomesial projection; propodus thick. P-3, P-4 Fungicola utinomii.-Serene, 1966:396 [un coxae with well-developed anterior lobes. P-5 justified emendation]. dactylus rotated anteriorly. PLP-3 of female Pseudocryptochirus ishigakiensis Takeda & uniramous. Male with slender P-l; abdomen Tamura, 1979: 188 [TL; Ishigakijima Is length 1.2 times width, segment 3 widest, lat land , Ryukyu , Japan; holotype* NSMT-Cr eral margins markedly convex. PLP-l slightly 5898]. curved laterally, apex pointed; lateral margin Hiroia ishigakiensis.- Takeda & Tamura, with simple setae distally, pappose setae pro 1981b:20. ximally, mesial margin with pappose and sim ple setae. Fungicola fagei (Fize & Serene, 1956)

REMARKS: Because Fize and Serene (1957) Troglocarcinus fagei Fize & Serene, 1956a: did not designate a type species for Troglocar 378 [TL: Nhatrang, Vietnam ; location of cinus (Fungicola), the name Fungicola was un- type unknown]. FIGURE 4. Fungico/a utinomi (Fize & Serene). Female (Palau): a, dorsal view ofcarapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;f, MXP-3; g, endopod ofMXP-I; h, MXL-l. Male (Palau): i, abdomen;), PLP . Scale: a, b = I. Omm; c, d, i = 0.9 mm; e,f= 0.3 mm; g = 0.2 mm; h,j = 0.1 mm. 428 PACIFIC SCIENCE, Volume 44, October 1990

Troglocarcinus (Fungicola) fagei.-Fize & Hapalocarcinus marsupialis Stimpson, 1859: Serene, 1957: 131. 412 [TL: Hilo, Hawaii ; location oftype un Fungicola fagei.- Serene, 1966: 397. known]. Cryptochirus marsupialis.- Taylor, 1971 : Genus Hapalo carcinus Stimpson, 1859 100. Hapalocarcinus Stimpson, 1859:412 [type Genus Hiroia Takeda & Tamura, 1981 species: Hapalocarcinus marsupialis Stimp son, 1859:412, by monotypy; gender mas Hiroia Takeda & Tamura, 1981b:20 [type culine]. species: Troglocarcinus krempfi Fize & Serene, 1956a:378, by original designation; DIAGNOS IS: Carapace quadrate, as long as gender feminine]. wide, widest at midlength, slightly convex in lateral view, not deflected anteriorly, lacking DIAGNOSIS: Carapace rectangular, longer depressions; mesogastric region not inflated. than broad, widest at midlength, flat in lateral Cardiointestinal region not outlined by de view, not deflected anteriorly, with broad, pressions. Pterygostomial region fused to ca shallow W-shaped depression; mesogastric re rapace. Epistome without lateral ridges, an gion slightly inflated. Cardiointestinal region terior margin sinuous with deep, wide median outlined by depression. Pterygostomial region indentation. Antennule not produced past fused to carapace. Epistome with subparallel midlength of eyestalk, apex pointed or lateral ridges, converging slightly anteriorly, rounded. Antennal segment 2 longer than anterior margin sinuous with slight median broad, margin lacking lateral spine. MXP-3 indentation. Lateral lobe of antennule oval, with exopod; merus without distolateral pro extending beyond eyestalk. Antennal segment jection, mesial margin lacking setae. Second 2 longer than broad, distal margin lacking segment of MXP-2 endopod with setae at lateral spine. MXP-3 with exopod; merus with distal margin of inner surface. MXP-I endo distolateral projection, mesial margin with pod quadrangular. Ventral thorax wider than setae. Inner surface of second segment of long, flat. Sternite of P-l smooth. Sternite of MXP-2 endopod with setae at distal margin. P-4 without suture. Femal e gonopore circu MXP-I endopod triangular, widest distal to lar, without hood. P-I dactylus with or with midlength; distal margin convex. Ventral out proximal tooth on cutting edge. P-2 merus thorax longer than wide, concave mesially. lacking distomesial projection, propodus Anterior extension ofsternite of P-I with few thin. P-3, P-4 coxae lacking anterior lobes. P-5 granules. Sternite of P-4 with suture. Female dactylus not rotated anter iorly. PLP-3 of fe gonopore oval, with anterior hood. P-I cut male uniramous. Ma le with robust P-I ; abdo ting edges entire. P-2 merus lacking disto men length 1.7 times width, segment 4 widest, mesial projection; propodus thick. P-3 coxa lateral margins convex. PLP-I curved later lacking, P-4 coxa with reduced an terior lobe. ally, apex pointed, lateral margin lacking P-5 dactylus rotated anteriorly. PLP-3 of fe setae, mesial margin with few pappose setae. male uniramous. Male with robust P-I ; abdo men length 1.9 times width, segment 4 widest, HOSTS: Pocilloporidae- Pocillopora, Stylo lateral margins con vex.PLP-I slightly curved phora, Seriatopora (see Fize and Serene 1957). laterally, apex pointed, lateral margin with DEPTH: < I to 27 m (RKK). several pappose, with few simple setae, mesial margin with several simple setae proximally. DISTRIB UTION: Indo-West Pacific-South Africa and Red Sea to Colombia (see refer HOSTS: Faviidae-Cyphastrea, Hydno ences in Takeda and Tamura 1986). phora (see Fize and Serene 1957, RKK). Merulinidae-Merulina (see Fize and Serene 1957). Hapalocarcinus marsupialis Stimpson, 1859 Figure 5 DEPTH: I to 19 m (RKK). FIGURE5. Hapalocarcinus marsupialis Stimpson. Female (Gu am): a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistom e; f, MXP-3; g, endopod of MXP-l; h, MXL-1. Male (Papu a New Gu inea): i, abdomen;j, PLP (from MacNamee 1961). Scale: a- c = 2.1 mm; d = 1.0 mm; i = 0.4 mm. j"= 0.3 mm; e. g = 0.2 mm; h = 0.1 mm;j, not to scale.

DISTRIBUTION: Vietnam (Fize and Serene 378 [TL: Nhatrang, Vietnam ; location of 1957); Japan-Izu Islands, Ryukyu Islands type unknown] . (Takeda and Tamura 1981b); Micronesia Troglocarcinus (Troglocarcinus) krempji. Palau, Guam (RKK). Fize & Serene, 1957: 79. Pseudocryptochirus krempfi.c-Seiene, 1966: Hiroia krempji (Fize & Serene, 1956) 396. Figure 6 Hiroia krempji.- Takeda & Tam ura, 1981b: 20. Troglocarcinus krempji Fize & Serene, 1956a: 430 PACIFIC SCIENCE, Volume 44, October 1990

FIGURE 6. Hiroia krempfi (Fize & Serene). Female (Palau) : a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;f, MXP-3; g, endopod ofMXP-I; h, MXL-1. Male (Pohnpei) : i, abdomen;}, PLP . Scale: a- c = 1.0 mm; d, i = 0.5 mm;f= 0.3 mm; e = 0.2 mm; g, h.] = 0.1 mm. Revision of Pacific Gall Crab Genera -s-Knorr 431

Genus Lithoscaptus Milne Edwards, 1862 Palau, Guam, Pohnpei (RKK); Palmyra Is land and Teraina (Edmondson 1933). Lithoscaptus Milne Edwards, 1862: FlO [type species: Lithoscaptus paradoxus Milne Ed wards, 1862:FlO, by monotypy; gender Lithoscaptus paradoxus Milne Edwards, 1862 masculine]. Figure 7

DIAGNOSIS : Carapace rectangular, longer Lithoscaptus paradoxus Milne Edwards, than broad, widest near midlength, sharply 1862:FlO [TL: Reunion Island; lectotype* convex in lateral view, deflected anteriorly, MNHN]. with broad W-shaped depression; mesogastric Cryptochirus coralliodytes var. rubrolineata region inflated . Cardiointestinal region out Fize & Serene, 1957:40 [TL: Nhatrang, lined by inverted If-shaped depression. Ptery Vietnam; location of type unknown]. gostomial region fused to carapace. Epistome Cryptochirus coralliodytes var. fusca Fize & with parallel lateral ridges, anterior margin Serene, 1957: 40 [TL: Nhatrang, Vietnam; straight with slight median indentation. Lat location of type unknown]. erallobe of antennule oval, extending beyond Cryptochirus coralliodytes var. parvulus Fize eyestalk. Antennal segment 2 longer than & Serene, 1957:40 [TL: Nhatrang, Viet broad, distal margin lacking lateral spine. nam; location of type unknown]. MXP-3 with exopod; merus with distolateral Cryptochirus bani Fize & Serene, 1957: 44 projection, mesial margin with setae. Inner [TL: Nhatrang, Vietnam; location of type surface ofsecond segment ofMXP-2 endopod unknown]. with setae at distal margin. MXP-l endopod triangular, widest distal to midlength; distal ?Lithoscaptus pacificus (Edmondson, 1933), margin convex. Ventral thorax longer than new combination wide, concave mesially. Anterior extension of sternite of P-I smooth. Sternite of P-4 with Cryptochirus pacificus Edmondson, 1933: 8 suture. Female gonopore oval, with anterior [TL: Palmyra Island; holotype* BPBM hood. P-I dactylus cutting edge with tooth S3669]. proximally. P-2 merus lacking distomesial REMARKS: The type of Cryptochirus paci projection; propodus slender. P-3, P-4 coxae ficu s Edmondson is not in good condition; with reduced anterior lobes. P-5 dactylus ro therefore the placement of that species in tated anteriorly. PLP-3 of female uniramous. Lithoscaptus is tentative. Male with robust P-l; abdomen length 3.8 times width, segments subequal in width, lat Lithoscaptus tri (Fize & Serene, 1956), new eral margins parallel. PLP-I curved laterally, combination apex pointed; lateral margin with simple setae, mesial margin with shorter simple setae Crytochirus tri Fize & Serene, 1956b: 379 [TL: proximally. Nhatrang, Vietnam; location of type un known] . HOSTS: Echino Faviidae-Cyphastrea, Cryptochirus trii.-Takeda & Tamura, pora, Favia, Favites, Hydnophora, Goniastrea, 1980d: 51 [unjustified emendation]. Leptastrea, Platygyra , Plesiastrea (see Fize and Serene 1957, RKK). Merulinidae REMARKS: The emendation of the specific M erulina (see Fize and Serene 1957). name tri to trii by Takeda and Tamura (l980d) was not justified, as the root of the DEPTH: < I to 12 m (RKK). name is "tr" (see analogous argument for DISTRIBUTION: Reunion (Milne Edwards Fungicola utinomi in " Remarks" section under 1862); Vietnam (Fize and Serene 1957); the genus Fungicola) . Japan-Izu Islands, Kushimoto, Ogasawara Islands, Ryukyu Islands (Takeda and Tamura Lithoscaptushelleri (Fize & Serene, 1957), new I980d, 1981a, 1983, 1986); Micronesia- combination 432 PACIFIC SCIENCE, Volume 44, October 1990

FIGURE 7. Lithoscaptus paradoxus Milne Edwards. Female (Guam): a, dorsal view ofcarapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;f, MXP-3; g, endopod of MXP-1. Male (Guam): h, abdomen; i, PLP . Scale: b = 2.1 mm; c = 1.0 mm; a = 0.8 mm; d, h = 0.7 mm;f= 0.5 mm; e = 0.3 mm; g = 0.2 mm; i = 0.1 mm. Revision of Pacific Gall Crab Genera-KROPP 433

Troglocarcinus (Favicola) helleri Fize & Serene, circular, with anterior hood. P-I cutting edges 1957:93 [TL: Nhatrang, Vietnam; location entire. P-2 merus with distomesial projection; of type unknown]. propodus thick. P-3, P-4 coxae with well Favicola helleri.c-Seteo», 1966: 397. developed anterior lobes. P-5 dactylus not ro tated anteriorly. PLP-3 of female biramous. Lithoscaptus nami (Fize & Serene, 1957), new Male with slender P-l; abdomen length 1.7 combination times width, segment 3 widest, lateral margins convex. PLP-I angularly curved laterally, Cryptochirus nami Fize & Serene, 1957:46 apex blunt, lateral and mesial margins with [TL: Nhatrang, Vietnam; location of type long pappose setae at midlength. unknown]. REMARKS: Fize and Serene (1957) did not Lithoscaptus grandis (Takeda & Tamura, designate a type species for Neotroglocarcinus, 1983), new combination rendering that name unavailable (ICZN Art. 13(b)]. Takeda and Tamura (1980c), by desig Cryptochirus grandis Takeda & Tamura, nating Troglocarcinus monodi as the type spe 1983: 2 [TL: Kushimoto, Japan; holotype* cies of the genus, made the name Neotroglo NSMT-Cr 8551]. carcinus available. Kropp (1988a) determined T. monodi to be a subjective junior synonym Genus Neotroglocarcinus Takeda & Tamura, of Cryptochirus hongkongensis Shen. 1980 HOSTS: Dendrophyllidae-Turbinaria (see Neotroglocarcinus Fize & Serene, 1957: 135 Fize and Serene 1957). [name unavailable]. DEPTH: < I to 13 m (RKK). Neotroglocarcinus Takeda & Tamura, 1980c: 147 [type species: Troglocarcinus monodi DISTRIBUTION: Vietnam (Fize and Serene Fize & Serene, 1956a: 375, a subjectivejun 1957); Japan-Izu Islands, Ryukyu Is ior synonym ofCryptochirus hongkongensis lands (Takeda and Tamura 1980c, 1986); Shen, 1936 (see Kropp 1988a), by original Micronesia-Palau, Guam, Pohnpei designation; gender masculine]. (RKK), Enewetak (Garth et al. 1987). Neotroglocarcinus hongkongensis (Shen, 1936) DIAGNOSIS: Carapace vase-shaped, longer than broad, widest posterior to midlength, flat Figure 8 in lateral view, deflected anteriorly; with irre Cryptochirus hongkongensis Shen, 1936: 23 gular depression flanking mesogastric region, [TL: Hong Kong; location of type un latter slightly inflated. Cardiointestinal region known]. outlined by depression. Pterygostomial region Pseudocryptochirus hongkongensis.- Takeda not fused to carapace. Epistome with parallel & Tamura, 1981b: 15. lateral ridges, anterior margin sinuous with Troglocarcinus monodi Fize & Serene, 1956a: deep, narrow median indentation. Lateral 375 [TL: Nhatrang, Vietnam; location of lobe of antennule triangular, extending be type unknown]. yond eyestalk. Antennal segment 2 longer Neotroglocarcinus monodi.-Fize & Serene, than broad, distal margin lacking lateral 1957: 137. spine. MXP-3 with exopod; merus without Neotroglocarcinus hongkongensis.- Kropp, distolateral projection, mesial margin with 1988a: 870. setae. Inner surface of second segment of MXP-2 endopod lacking setae at distal mar Neotroglocarcinus dawydofji (Fize & Serene, gin. MXP-I endopod triangular, widest near 1956) distal margin; latter concave. Ventral thorax about as long as wide, flat. Anterior extension Troglocarcinus dawydofji Fize & Serene, of sternite of P-I with tubercles. Sternite of 1956a :376 [TL: Nhatrang, Vietnam; loca P-4 lacking suture. Female gonopore semi- tion of type unknown]. 434 PACIFIC SCIENCE, Volume 44, October 1990

FIGURE 8. Neotroglocarcinus hongkongensis (Shen). Female (Vietnam): a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;f, MXP-3; g, endopod of MXP-I ; h, MXL-I . Male (Vietnam): i, abdomen;j, PLP. Scale: a-c = 2.1 mm; d = 1.0 mm; e = 0.5 mm;f= 0.2 mm; g, h,j = 0.1 mm. Revision of Pacific Gall Crab Genera-e-Knor-e 435

Neotroglocarcinus dawydoffi .-Fize & Se the west coast of Mexico (see Kropp 1989). rene, 1957: 144. Also Atlantic (Kropp and Manning 1987).

Opecarcinus crescentus (Edmondson, 1925) Genus Opecarcinus Kropp & Manning, 1987 Cryptochirus crescentus Edmondson, 1925: 33 Opecarcinus Kropp & Manning, 1987:9 [type [TL: Johnston Island; holotype* BPBM species: Pseudocryptochirus hypostegus SI805]. Shaw & Hopkins, 1977: 179, by original Pseudocryptochirus crescentus.-Utinomi, designation; gender masculine]. 1944: 697. DIAGNOSIS: Carapace vase-shaped, longer Troglocarcinus (Troglocarcinus) crescentus. than broad, widest posterior to midlength, Fize & Serene, 1957:62. con vex in lateral view, deflected anteriorly, Opecarcinus crescentus.-Kropp & Manning, with transverse depression on protogastric re 1987:9 ; Kropp 1989:99. gion; mesogastric region slightly inflated . Cardiointestinal region outlined by depres Opecarcinus granulatus (Shen, 1936) sion. Pterygostomial region fused to carapace. Cryptochirus granulatus Shen, 1936: 23 [TL: Epistome with paraIlellateral ridges, anterior Hong Kong; holotype* BMNH 1911 .8.15.2]. margin sinuous with shaIlow, wide median Opecarcinus granulatus. -Kropp, 1989: 104. indentation. Lateral lobe of antennule trian gular, extending beyond eyestalk. Antennal Opecarcinus hypostegus (Shaw & Hopkins, segment 2 longer than broad, distal margin 1977) lacking lateral spine. MXP-3 with exopod; Figure 9 merus with distolateral projection, mesial Pseudocryptochirus hypostegus Shaw & Hop margin with setae. Inner surface of second kins, 1977: 179 [TL: Florida Middle segment of MXP-2 endopod lacking setae at Ground, Florida; holotype* NMNH distal margin. MXP-I endopod triangular, 168533]. widest near midlength; distal margin convex. Opecarcinus hypostegus.-Kropp & Man Ventral thorax longer than wide, flat. Anter ning, 1987: 10. ior extension ofsternite ofP-I with tubercles. Sternite of P-4 with suture. Female gonopore Opecarcinus aurantius Kropp, 1989 oval, lacking or with anterior hood. P-l cut ting edges entire. P-2 merus with distomesial Opecarcinus aurantius Kropp, 1989: 108 [TL: projection; propodus thick. P-3, P-4 coxae Cetti Bay, Guam; holotype* NMNH with well-developed anterior lobes. P-5 dac 234266]. tylus rotated anteriorly. PLP-3 of female uni Opecarcinus lobifrons Kropp, 1989 ramous. Male with robust P-I; abdomen length 1.8 times width , segment 3 widest, lat Troglocarcinus (Troglocarcinus) crescentus. eral margins convex. PLP-I curved lateraIly , Garth, 1965: 8 [not Cryptochirus crescentus apex blunt, lateral margin with several, mesial Edmondson]. margin with few stout simple setae. Pseudocryptochirus crescentus.-Garth & Hopkins, 1968:41 [in part; not Crypto HOSTS: Agariciidae- Agaricia, Gardinero seris, Leptoseris, Pavona (see Fize & Serene chirus crescentus Edmondson]. 1957, Scott 1985, RKK). Siderasteriidae Opecarcinus lobifrons Kropp, 1989: 112 (TL: Siderastrea (see Scott 1985), Coscinaraea Tanguisson Point, Guam; holotype* [recorded as ?Coscinastrea of Agariciidae by NMNH 234269]. Takeda and Tamura (1983)]. Opecarcinus peliops Kropp, 1989 DEPTH: < I to 82 m (Shen 1936, RKK). Opecarcinus peliops Kropp, 1989: 116 [TL: DISTRIBUTION: Christmas Island (Indian Pakin Atoll, Pohnpei; holotype* NMNH Ocean) (Shen 1936); Vietnam and Japan to 234272]. 436 PACIFIC SCIENCE, Volume 44, October 1990

FIGURE 9. Opecarcinus hypostegus (Shaw & Hopkins). Female holotype (Florida): a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome ; f, MXP-3; g, endopod of MXP-I ; h, MXL -1. Male paratype (Florida): i, abdomen; i, PLP. Scale: c = 1.2 mm; b, d = 1.0 mm; a = 0.7 mm; f, i = 0.5 mm; e = 0.3 mm; h,j = 0.2 mm; g = 0.1 mm. Revision ofPacific Gall Crab Genera-KRopp 437

Opecarcinus pholeter Kropp, 1989 crab, in reference to the bowl-shaped conca vities on the anterior carapace. Opecarcinus pholeter Kropp, 1989: 118 [TL: Cetti Bay, Guam; holotype* NMNH REMARKS: Pelycomaia is most closely re 234275]. lated to Fungicola (see Kropp 1988d), but can be distinguished from it by the two bowl Opecarcinus sierra Kropp, 1989 shaped concavities on the anterior carapace and by having a reduced anterior lobe on the pecarcinus sierra Kropp, 1989: 122 [TL: o coxa ofP-3. Fungicola lacks bowl-shaped con Agat Bay, Guam; holotype* NHNH cavities on the carapace and has a well 234278]. developed lobe on the coxa of P-3. HOSTS: Faviidae -Cyphastrea, Leptastrea Genus Pelycomaia Kropp, new genus (see Fize & Serene 1957). Pelycomaia Kropp, new genus [type species DEPTH: <2 m (RKK). Cryptochirus minutus Edmondson, 1933: 12, by original designation; gender DISTRIBUTION: Vietnam (Fize & Serene feminine]. 1957); Micronesia-Guam (RKK); Hawai'i (Edmondson 1933). DIAGNOSIS: Carapace rectangular, longer than broad, widest anterior to midlength, con Pelycomaia minuta (Edmondson, 1933) vex in lateral view, deflected anteriorly, with Figure 10 two bowl-shaped concavities anteriorly; me sogastric region slightly inflated. Cardiointes Cryptochirus minutus Edmondson, 1933: 12 tinal region outlined by depression. Pterygo [TL: Oahu, Hawaii, holotype* BPBM stomial region fused to carapace. Epistome S3671]. with parallel lateral ridges, anterior margin Troglocarcinus (Favicola) minutus.-Fize & straight with slight median indentation. Lat Serene, 1957: 106. eral lobe of antennule oval, extending just Favicola minutus.-Serene, 1966: 397. beyond eyestalk. Antennal segment 2 longer Favicola minutum.- Serene et al., 1976: 20. than broad, distal margin lacking lateral Favicola minuta. -Takeda & Tamura, 1981 a : spine. MXP-3 with exopod; merus with disto 43. lateral projection, mesial margin with setae. Inner surface of second segment of MXP-2 Genus Pseudocryptochirus Hiro, 1938 endopod with setae at distal margin. MXP-l endopod triangular, widest to midlength; Pseudocryptochirus Hiro, 1938: 149[type spe distal margin convex. Ventral thorax longer cies: Pseudocryptochirus viridis Hiro, 1938: than wide, concave mesially. Anterior exten 149, by monotypy; gender masculine]. sion of sternite of P-l smooth. Sternite of P-4 DIAGNOSIS: Carapace vase-shaped, longer with suture. Female gonopore elliptical, with than broad, widest posterior to midlength, anterior hood. P-l dactylus with low tooth convex in lateral view, not deflected anter proximally on cutting edge. P-2 merus lacking iorly, lacking depression; mesogastric region distomesial projection; propodus thick. P-3 very slightly inflated. Cardiointestinal region coxa with reduced , P-4 coxa with well not outlined by depression. Pterygostomial developed anterior lobes. P-5 dactylus rotated region not fused to carapace. Epistome with anteriorly. PLP-3 of female uniramous. Male parallel lateral ridges, anterior margin sinuous with robust P-l; abdomen length 3.0 times with shallow, narrow median indentation. width , segment 5 widest, lateral margins sub Lateral lobe of antennule quadrangular, ex parallel. PLP-l not available for study. tending beyond eyestalk. Antennal segment 2 ETYMOLOGY: From the Greek, pelykos, shorter than broad, distal margin lacking lat bowl, in combination with maia, a kind of eral spine. MXP-3 with exopod; merus with- 438 PACIFIC SCIENCE, Volume 44, October 1990

FIGURE 10. Pelycomaia minuta (Edmondson). Female (Hawai'i): a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;f, MXP- 3; g, endopod of MXP-I ; h, MXL-I . Male (Hawa i'i, from Edmond son 1933): i, abdomen. Scale: c = 0.8 mm; a, b, d = 0.5 mm;!= 0.2 mm; e, g, h = 0.1 mm; i, not to scale. out distolateral projection, mesial margin of P-4 with partial suture. Female gonopore with setae. Inner surface ofsecond segment of oval, with anterior hood. P-I cutting edges MXP-2 endopod lacking setae at distal mar entire. P-2 merus with distomesial projection; gin. MXP-I endopod triangular, widest near propodus thick . P-3, P-4 coxae with well distal margin ; latter concave. Ventral thorax developed anterior lobes. P-5 dactylus not about as long as wide, flat. Anterior extension rotated anteriorly. PLP-3 of female uniram of sternite of P-l with few granules. Sternite ous. Male with slender P-I ; abdomen length Revision of Pacific Gall Crab Genera-c-Kaozr 439

FIGURE II. Pseudocryptochirus viridis Hiro . Female (Palau) : a, dorsal view of carapace ; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;j, MXP-3; g, endopod of MXP-I; h, MXL-1. Male (Palau): i, abdomen ;j, PLP. Scale: a = 1.2mm; b, c = 1.0 mm; d = 0.5 mm; e.f= 0.2 mm; g, h,j= 0.1 mm.

1.4 times width, segment 3 widest, lateral 1957); Indonesia (Serene et al. 1974); Japan margins concave. PLP-I slightly curved late Izu Islands (Takeda and Tamura 1981b, rally, apex sharply pointed, lateral and mesial 1986); Micronesia-Palau, Guam, Pohnpei margins with long pappose setae at midlength. (RKK). A record of the species from Enewe HOSTS: Dendrophylliidae-Turbinaria (see tak (Garth and Hopkins 1968) is apparently Fize & Serene 1957). erroneous (Garth et al. 1987). DEPTH: I to 6 m (RKK). Pseudocryptochirus viridis Hiro, 1938 DISTRIBUTION: Vietnam (Fize & Serene Figure II 440 PACIFIC SCIENCE, Volume 44, October 1990

Pseudo cryptochirus viridis Hiro, 1938: 150 Pseudohapalocarcinus ransoni Fize & Serene, [TL: Tanabe Bay, Japan; location of type 1956 unknown]. Figure 12 Pseudohapalocarcinus ransoni Fize & Serene, Genus Pseudohapalocarcinus Fize & Serene, 1956a: 378 [TL: Nhatrang, Vietnam; loca 1956 tion of type unknown]. Pseudohapalocarcinus Fize & Serene, 1956a: 378 [type species: Pseudohapalocarcinus Genus Sph enomaia Kropp, new genus ransoni Fize & Serene, 1956a: 378, by mo Sphenomaia Kropp, new genus [type species: notypy; gender masculine]. Cryptochirus pyriformis Edmondson, 1933: DIAGNOSIS : Carapace subquadrate, as long 10, by original designation and monotypy; as broad, widest posterior to midlength, gender feminine]. evenly convex in lateral view, not deflected DIAGNOSIS: Carapace rectangular, longer anteriorly, lacking depression; mesogastric re than broad, widest at midlength, strongly con gion not inflated. Cardiointestinal region not vex in lateral view, not deflected anteriorly, outlined by depression. Pterygostomial region with broad depression over entire anterior fused to carapace. Epistome with parallellat portion; mesogastric region not inflated. Car eraI ridges, anterior margin sinuous with diointestinal region outlined by inverted U deep, wide median indentation. Lateral lobe shaped depression. Pterygostomial region of antennule quadrangular, not extending be fused to carapace. Epistome with parallel lat yond eyestalk. Antennal segment 2 shorter eral ridges, anterior margin sinuous with shal than broad, distal margin lacking lateral low, wide median indentation. Lateral lobe of spine. MXP-3 with exopod; merus without antennule subquadrangular, extending to or distolateral projection, mesial margin with just beyond eyestalk . Antennal segment 2 setae. Inner surface of second segment of longer than broad, distal margin lacking lat MXP-2 endopod lacking setae at distal mar eral spine. MXP-3 with exopod; merus without gin. MXP-I endopod triangular, widest near distolateral projection, mesial margin with midlength; distal margin convex. Ventral setae. Inner surface of second segment of thorax wider than long, flat. Anterior exten MXP-2 endopod with setae at distal margin. sion ofsternite of P-I smooth. Sternite of P-4 MXP-I endopod triangular, widest distall y; without suture. Female gonopore semicir distal margin slightly convex. Ventral thorax cular, lacking anterior hood. P-l cutting edges longer than wide, concave mesially. Anterior entire. P-2 merus lacking distomesial projec extension of sternite of P-I with granules. tion ; propodus slender. P-3, P-4 coxae with Sternite of P-4 with suture. Female gonopore well-developed anterior lobes. P-5 dactyl us elliptical , with anterior hood. P-I cutting not rotated anteriorly. PLP-3 offemale unira edges entire. P-2 merus lacking distomesial mous. Male with robust P-I ; abdomen length projection; propodus slender. P-3, P-4 coxae 1.4times width, segment 4 widest, lateral mar with well-developed anterior lobes. P-5 dac gins convex. PLP-I slightly curved distally, tylus rotated anteriorly. PLP-3 of female uni apex blunt, lateral and mesial margins with ramous. Male not available for study. few simple setae. ETYMOLOGY: From the Greek, sphenos, HOSTS: & Agariciidae-Pavona (see Fize wedge, in combination with maia , a kind of Serene 1957). crab, in reference to the appearance of the car DEPTH: < I to 21 m (RKK). apace in lateral view.

DISTRIBUTION: Vietnam (Fize & Serene REMARKS: Sphenomaia is most closely re 1957); Japan-Ryukyu Islands (Takeda and lated to a group of genera: Cryptochirus, Tamura 1980a); Micronesia-Palau, Guam, Lithoscaptus, Fizesereneia , and Xynomaia (see Pohnpei (RKK). Kropp 1988d). Sphenomaia, Cryptochirus, Revision of Pacific Gall Crab Genera-e-Knor-e 441

e )-~\

FIGURE 12. Pseudohapalocarcinus ransoni (Fize & Serene). Female (Guam): a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;J, MXP-3; g, endopod ofMXP-I; h, MXL-1. Male (Palau): i, abdomen; j , PLP . Scale: a. b = 1.0 mm; c. d. i = 0.5 mm; e.f= 0.3 mm; g. h.] = 0.1 mm.

and Lithoscaptus, by having the pterygosto HOSTS: Faviidae-Favites, Hydnophora , mial region fused to the carapace, differ from Platygyra (see Edmondson 1933). Fizesereneia and Xynomaia, in which the DEPTH: Not recorded. pterygostomial region is not fused to the car apace. The clusters of rounded tubercles and DISTRIBUTION: Central Pacific- Teraina network ofgrooves on the posterior carapace (Edmondson 1933). of Cryptochirus distinguish it from Spheno maia, which has isolated tubercles and lacks Sphenomaia pyriforma (Edmondson, 1933), grooves on the posterior carapace. Spheno new combination maia, by having a sinuous anterior margin of Figure 13 the epistome, differs from Lithoscaptus, which Cryptochirus pyriformis Edmondson, 1933: has a straight anterior margin. 10 [TL: Washington Island (now known 442 PACIFIC SCIENCE, Volume 44, October 1990

FIGURE 13. Sphenomaia pyriforma (Edmondson). Female (Teraina) : a, dorsal view of cara pace; b, lateral view; c, thoracic sternites; d, P-2; e, episto me; f, MXP-3; g, endo pod of MXP-I. Scale: a-d = 1.0 mm; e = 0.5 mm: 1= 0.3 mm; g = 0.2 mm.

as Teraina, Kiribati); holotype* BPBM in lateral view, not deflected anteriorly, with S3670]. broad, shallow depre ssion; mesogastric re Troglocarcinus (Favicola) pyrijormis.-Fize gion slightly inflated. Cardiointestinal region & Serene, 1957:84. outlined by depression. Pterygostomial region Favicola pyrijormis.-Serene, 1966: 397. fused to carapace. Epistome lacking lateral ridges, ant erior margin sinuous with shallow, wide median indentation. Lateral lobe of Genus Utinomiella Kropp & Takeda, 1988 antennule rectangular, extending to tip ofeye Utinomiella Kropp & Takeda, 1988: 29 [type stalk. Antennal segment 2 longer than broad, species: Cryptochirus dimorphus Hender distal margin lacking lateral spine. MXP-3 son, 1906:214, by original designation and lacking exopod; merus without distol ateral monotypy; gender feminine]. projection, mesial margin without setae. Inner Utinomia Takeda & Tamura, 1981b:23 [aju surface ofsecond segment ofMXP-2 endopod nior homonym of Utinomia Tomlinson, with setae at distal margin. MXP-I endopod 1963:264 (Crustacea: Acro thoracica)]. triangular, widest near distal margin; latter convex. Ventral thorax longer than wide, flat. DIAGNOSIS : Carapace vase-shaped, longer Anterior extension of sternite of P-I smooth. than broad, widest posterior to midlength, flat Sternite of P-4 with partial suture. Female Revision of Pacific Gall Crab Genera-e-Kaor-r 443 gonopore semicircular, lacking anterior hood. parallel lateral ridges, anterior margin P-l cutting edges entire . P-2 merus lacking straight with slight median indentation. Lat distomesial projection; propodus slender. eral lobe of antennule oval, extending to tip P-3 coxa with reduced, P-4 coxa with well of eyestalk. Antennal segment 2 longer than developed anterior lobes. P-5 dactylus not ro broad, distal margin lacking lateral spine. tated anteriorly. PLP-3 of female uniramous. MXP-3 with exopod; merus with distolateral Male with robust P-l; abdomen length 1.7 projection, mesial margin with setae. Inner times width, segment 3 widest, lateral margins surface of second segment ofMXP-2 endopod convex. PLP-I slightly curved distally, apex with setae at distal margin. MXP-l endopod blunt, lateral and mesial margins with few triangular, widest proximal to midlength; long pappose setae distally. distal margin evenly convex. Ventral thorax longer than wide, flat. Anterior extension of HOSTS: Pocilloporidae-Pocillopora, Stylo phora (see Takeda and Tamura 1981b, RKK). sternite of P-I smooth or with fine granules. Sternite of P-4 with suture. Female gonopore DEPTH: 1 to 29 m (RKK). elliptical, with anterior hood. P-I dactylus with low tooth on cutting edge. P-2 merus lacking DISTRIBUTION: Andaman Islands (Hender son 1906); Japan-Ryukyu Islands (Takeda distomesial projection; propodus slender. P-3 and Tamura 1981b; Micronesia-Palau, coxa with poorly developed , P-4 coxa with Guam, Pohnpei (RKK); Hawai'i (McCain well-developed anterior lobe. P-5 dactylus ro and Coles 1979). tated anteriorly. PLP-3 of female uniramous. Male with robust P-I ; abdomen length 2.8 times width , segment 4 widest, lateral margins Utinomiella dimorpha (Henderson, 1906) subparallel. PLP-l slightly curved distally, Figure 14 apex pointed, lateral margin with many proxi Cryptochirus dimorphus Henderson, 1906:214 mally pappose setae, few simple setae, mesial [TL: Andaman Islands ; location oftype un margin with several short simple setae. known]. ETYMOLOGY: From the Greek, xynon , com Pseudocryptochirus kahe McCain & Coles, panion, in combination with maia, a kind of 1979:81 [TL: Kahe Point, O'ahu, Hawai 'i; crab, in reference to the symbiotic life-style of holotype* BPBM S8512]. the crab. Pseudocrypto chirus dimorphus.-Takeda & Tamura, 1980d:47. REMARKS: Xynomaia is most closely related Utinomia dimorpha.- Takeda & Tamura, to Fizesereneia (see Kropp 1988d), but is dis 1981b:23. tinguished from it by having a W-shaped de Utinomiella dimorpha.-Kropp & Takeda, pression on the anterior carapace, not the two 1988: 29. bowl-shaped concavities on the anterior car apace characteristic of Fizesereneia. Also, the carapace of Xynomaia is not deflected anter Genus Xynomaia Kropp, new genus iorly, whereas that ofFizesereneia is deflected.

Xynomaia Kropp, new genus [type species: HOSTS: Faviidae-Favia, Goniastrea, Mon Troglocarcinus sheni Fize & Serene, 1956b: tastrea, Oulophyllia, Platygyra, (see Fize and 380, by original designation; gender femi Serene 1957, RKK). Merulinidae-Merulina nine]. (see Fize and Serene 1957). Pectiniidae DIAGNOSIS: Carapace rectangular, longer Pectinia (RKK). than broad, widest at midlength, convex in DEPTH: 1 to 15 m (RKK). lateral view, not deflected anteriorly, with broadly W-shaped depression; mesogastric DISTRIBUTION: Vietnam (Fize and Serene region slightly inflated. Cardiointestinal re 1957);Sumatra (Serene 1966); Japan-Izu Is gion outlined by depression. Pterygostomial lands, Kushimoto (Takeda and Tamura 1983, region not fused to carapace. Epistome with 1986); Micronesia-Palau, Guam (RKK). 444 PACIFIC SCIENCE, Volume 44, October 1990

() n

~ h~ 9 :=:::s:::::-:_...... _ f ~

FIGURE 14. Utinomiella dimorpha (Henderson). Female (Palau): a, dorsal view of carapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;f, MXP-3; g, endopod ofMXP-I; h, MXL-l. Male (Guam) : i, abdomen ;}, PLP. Scale: a-c, i = 1.0 mm; d = 0.5 mm;f= 0.3 mm; e = 0.2 mm; g. h,j = 0.1 mm. Revision of Pacific Gall Crab Genera-KRopp 445

j gr

FIGURE 15. Xynomaia sheni (Fize & Serene). Female (Guam): a, dorsal view ofcarapace; b, lateral view; c, thoracic sternites; d, P-2; e, epistome;f, MXP-3 ; g, endopod of MXP-I ; h, MXL-1. Male (Guam): i, abdomen;j, PLP. Scale: a-c, i = 1.0 mm; e = 0.5 mm; d.f= 0.3 mm; g, h,j = 0.1 mm. 446 PACIFIC SCIENCE, Volume 44, October 1990

Xynomaia sheni (Fize & Serene, 1956), new LITERATURE CITED combination Figure 15 CALMAN, W. T. 1900. On a collection of Brachyura from Torres Straits. Trans. Troglocarcinus sheni Fize & Serene, 1956b:380 Linn. Soc. London, 2d Ser. (Zool.) 8(1): 1 [TL: Nhatrang, Vietnam; location of type 50, pI. 1-3. unknown]. CO~LHO, P. A. and M. A. RAMOS. 1972. A Troglocarcinus (Troglocarcinus) sheni.-Fize constituicao e a distribuicao de fauna de & Serene, 1957: 74. decapodos do litoral leste da America do Pseudocryptochirus sheni.-Serene, 1966: Sol entre las latitudes 5° N 39° S. Trab. 396. Oceanogr. Univ. Fed . Pernambuco 13: Hiroia sheni.-Takeda & Tamura, 1981b: 20. 133-236, figs. 1-4. EDMONDSON, C. H. 1925. Marine zoology of Xynomaia boissoni (Fize & Serene, 1956), new the central Pacific: Crustacea. Bull. Bernice combination P. Bishop Mus. 27: 3-62. Troglocarcinus boissoni Fize & Serene, 1956a : --- . 1933. Cryptochirus of the central Pa 376 [TL: Nhatrang, Vietnam; location of cific: Occas. Pap. Bernice Pauai Bishop type unknown]. Mus. 10(5): 1-23, figs. 1-6, pI. 1-4. Troglocarcinus (Troglocarcinus) boissoni. FIZE, A., and R. SERENE. 1956a. Note preli Fize & Serene, 1957: 69. minaire sur huit especes nouvelles, dont une Pseudocryptochirus boissoni-s-Seren«, 1966: d'un genre nouveau, D'Hapalocarcinides, 396. Bull. Soc. Zool. Fr. 80(5-6): 375-378, figs. Hiroia boissoni.- Takeda & Tamura, 1981b: 1-2. 20. - - -. 1956b. Note preliminaire quatre es peces nouvelles D'Hapalocarcinides avec Xynomaia verrilli (Fize & Serene, 1957) new quelques remarques au sujet du Crypto combinaton chirusrugosus Edmonson [sic]. Bull. Soc. Zool. Fr. 80(5 -6): 379-382, fig. I. Troglocarcinus (Favicola) verrilli Fize & Se - - - . 1957. Les hapalocarcinides du Viet rene, 1957: 101 [TL: Nhatrang, Vietnam; Nam. Arch. Mus. Natl. Hist. Nat. (Paris), location of holotype unknown, paratype ser. 7, 5: 1-202, figs. 1-43, pI. 1-18. MNHN (not seen)]. GARTH, J. S. 1965. The brachyuran decapod Favicola verrilli.- Serene, 1966: 397. crustaceans of Clipperton Island. Proc. Calif. Acad. Sci., 4th ser., 33 : 1-46, figs. 1-26. ACKNOWLEDGMENTS GARTH, J. S., and T. S. HOPKINS. 1968. Pseudocryptochirus crescentus (Edmond I thank the University of Guam Marine son), a secondcrab ofthe corallicolous fam Laboratory and the Micronesian Mariculture ily Hapalocarcinidae (Crustacea, Deca Demonstration Center for supporting my poda) from the eastern Pacific. Bull. South fieldwork in Micronesia and the Division of Calif. Acad. Sci. 67: 40-48, figs. 1-2. Crustacea (NMNH) for supporting my sys GARTH, J. S., J. HAIG, and J. W. KNUDSEN. tematics studies. Specimens used in this study 1987. Crustacea, Decapoda (Brachyura were kindly provided by A. Crosnier and Anomura) of Enewetak Atoll. Pages (MNHN), R. Ingle, (BMNH), M. Takeda 235-261 in D . M. Devaney, E. S. Reese , B. (NSMT), B. Burch (BPBM), and V. Stagl L. Burch, and P. Helfrich, eds. The natural (NMW). R. H. Randall identified the coral history of Enewetak Atoll. Vol. II. Biogeo hos ts of crabs collected in Micronesia. R. B. graphy and systematics. U. S. Dep. Energy, Manning, L. G . Eldredge, and G. J. Vermeij Washington, D.C. reviewed the manuscript. HELLER, C. 1861. Synopsis der im rothen Revision of Pacific Gall Crab Genera- KROPP 447

Meere vorkommenden Crustaceen. Verh. Decapoda), non Tomlinson, 1963 (Crusta Zool.-Bot. Ges. Wien 11: 1-32. cea, Acrothoracica). Bull. Biogeogr. Soc. HENDERSON, J. R. 1906. On a new species of Jpn.43 :29. coral-infesting crab taken by the R.I.M.S. LUND¢ ER, S. 1974. A checklist of the marine " Investigator" at the Andaman Islands. Brachyura in the reference collection at Ann. Mag. Nat. Hist., ser. 7,18:211-219, PMBC, Thailand. Phuket Mar. BioI. Cent. pI. 8. Res. Bull. 4 :3-11. H IRO, F. 1938. A new coral-inhabiting crab, MCCAIN, J. c., and S. L. COLES. 1979. A new Pseudocryptochirus viridis gen. et sp. nov . species of crab (Brachyura, Hapalocarcini (Hapalocarcinidae, Brachyura). ZooI. dae) inhabiting pocilloporid corals in Ha Mag. , Tokyo 50 : 149-151. waii. Crustaceana (Leiden) 36: 81-89, KRopp, R. K . 1986. Feeding biology and figs. 1-3, pI. I mouthpart morphology of three species of MACNAMEE, C. G. 1961. Life history, mor coral gall crabs (Decapoda: Cryptochiri phology, habits and taxonomy of Hapalo dae) . J. Crust. BioI. 6: 377-384, figs. 1-2. carcinus marsupialis Stimpson (Arthro ---. 1988a. The status of Cryptochirus poda, Crustacea, Decapoda). M.S. thesis , hongkongensis Shen , 1936 (Brachyura: Uni versit y of Hawaii, Honolulu. Cryptochiridae). Proc. BioI. Soc. Wash. McNEILL, F. A. 1968. Crustacea, Decapoda 101 :866-871, figs. 1-2. & Stomatopoda. Scientific Reports, Great - -- . 1988b. The status of Cryptochirus Barrier Reef Expedition, 1928-29,7(1): 1 coralliody tes Heller and Lithoscaptus para 98, figs. 1-2, pI. 1-2. doxus Milne Edwards (Brachyura: Crypto MILNE EDWARDS, A. 1862. Faune carcinologi chiridae). Proc. BioI. Soc. Wash. 101: 872 que de l' ile de la Reunion, Annexe F de 882, figs. 1-6. I'ouvrage intitule: Notes sur I' ile de la - --. 1988c. Case 2636. Fizesereneia Ta Reunion par L. Maillard. Pages FI-FI6, keda and Tamura, 1980 (Crustacea, Deca pI. 17-19 in L. Maillard, Notes sur l' i Ie de poda): Proposed confirmation of Troglo la Reunion (Bourbon). Paris. carcinus heimi Fize & Serene, 1956 as the MONOD, TH., and R. SERENE. 1976. Parasitic, type species. Bull. Zool. Nomencl. 45: 262 commensal, and inquiline crustaceans col 263. lected during the Rumphius Expedition II. --- . 1988d. Biology and systematics of Osean. Indones. 6: 23-27. coral gall crabs (Crustacea: Cryptochiridae). NOBILl, G. 1906. Faune carcinologique de la Ph.D. diss., University of Maryland, Col Mer Rouge. Decapodes et Stomatopodes. lege Park. Ann. Sci. Nat. Zool. (a) 4: 1-347. - --. 1989. A revision ofthe Pacific species PAULSON, O. 1875. Podophthalmata and of gall crabs, genus Opecarcinus (Crusta Edriophthalmata (Cumacea). Studies on cea: Cryptochiridae). Bull. Mar. Sci. 45 : Crustacea ofthe Red Sea with notes regard 98-129. ing other seas, Part I. xiv + 144p., pI. 1-21. KROPP, R. K., and R. B. MANNING. 1985. Kiev. [Orginal in Russian. Reprinted 1961, Cryptochiridae, the correct name for the in English, with different pagination, by the family containing the gall crabs (Crustacea: Israel Program for Scientific Translations, Decapoda: Brachyura). Proc. BioI. Soc. Jerusalem.] Wa sh. 98 :954-955. RICHTERS, F. 1880. Decapoda. Pages 139 - -- . 1987. The Atlantic gall crabs, family 178, pI. 15-18 in K. Mobius, Beitrage zur Cryptochiridae (Crustacea: Decapoda: Meeresfauna der Insel Mauritius und der Brachyura). Smithson. Contrib. Zool. 462 : Seychelles. 1-21 , figs. 1-10. SCOTT, P. J. B. 1985. Aspects of living coral KROPP, R. K., and M. TAKEDA. 1988. Utino associates in Jamaica. Proc. 5th Int. Coral miella, a replacement name for Utinomia Reef Congr., Tahiti 5:345-350. Takeda and Tamura, 1981 (Crustacea, SERENE, R. 1966. Note sur la taxonomie et la 448 PACIFIC SCIENCE, Volume 44, October 1990

distribution geographic des Hapalocarcini - --. 1980c. Coral-inhabiting crabs of the dae (Decapoda-Brachyura). Proc. Symp. family Hapalocarcinidae from Japan. IV. Crustacea, Ernakulam, 12-15 January Genus Neotroglocarcinus. Bull. Natl. Sci. 1965, Mar. BioI. Assoc. India Part I: 395 Mus. Tokyo, ser. A (Zool.) 6(3): 147-151., 398. figs. 1-2. SERENE, R., K. ROMIMOHTARTO, and M. K. - -- .I980d. Coral-inhabiting crabs of the MOOSA. 1974.The Hippidea and Brachyura family Hapalocarcinidae from Japan. V. collected by the Rumphius Expedition I. Genus Cryptochirus. Res. Crustacea 10: Osean . Indones. 1: 17-26. 45-56, text-figs. 1-2, pI. 2-4. --- . 1976. Hippidea, Brachyura, and - -- . 1981a. Coral-inhabiting crabs of the Stomatopoda of the Rumphius Expedition family Hapalocarcinidae from Japan. VII. II. Osean. Indones. 6: 15-21. Genus Favicola. Res. Crustacea II: 41-50, SHAW, K., and T. S. HOPKINS. 1977. The dis text-figs. 1-3, pI. 1-3. tribution of the family Hapalocarcinidae - --. 1981b. Coral-inhabiting crabs of the (Decapoda, Brachyura) on the Florida family Hapalocarcinidae from Japan. VIII . Middle Ground with a description of Genus Pseudocryptochirus and two new Pseudocryptochirus hypostegus new species. genera . Bull. Biogeogr. Soc. Jpn 36(3): 14 Proc. 3rd Int. Coral Reef Symp., Miami 27,figs. 1-3,pl. 1-4. 1: 177-183, figs. 1-3. ---. 1983. Coral-inhabiting crabs of the SHEN, C. J. 1936. Notes on the family Hapalo family Hapalocarcinidae from Japan. IX. A carcinidae (coral-infesting crabs) with des small collection made at Kushimoto and criptions of two new species. Hong Kong Koza, the Kii Peninsula. Bull. Natl. Sci. Nat. Suppl. (5): 21-26, pI. 1-2. Mus. Tokyo, ser. A (Zool.) 9(1): 1-11 , figs. STIMPSON, W. 1859. [Communication, Hapa 1-4, pI. 2-4. locarcinus marsupialis.] Proc. Boston Soc. ---. 1986. Coral-inhabiting crabs of the Nat. Hist. 6:412-413. family Hapalocarcinidae from Japan. XI. TAKEDA, M., and Y. TAMURA. 1979. Coral Biogeographical distribution. Bull. Bio inhabiting crabs of the family Hapalocar geogr. Soc. Jpn 41(8): 61-70, figs. 1-9. cinidae from Japan. I. Three species ob TAYLOR, J. D. 1971. Crustacea: Brachyura tained from mushroom coral , Fungia. Bull. and Anomura from Diego Garcia. Atoll Natl. Sci. Mus. Tokyo, ser. A (Zool.) 5(3): Res. Bull. 149: 93-101. 183-192, figs. 1-4. TOMLINSON, J. T. 1963. Two new acrotho --- . 1980a. Coral-inhabiting crabs of the racican barnacles from Japan. Pub I. Seto family Hapalocarcinidae from Japan. II. Mar. BioI. Lab. 11 (2): 263-280. Genus Pseudohapalocarcinus. Proc. Jpn. UTINOMI , H. 1944. Studies on the animals in Soc. Syst. Zool. 18:54-59, pI. 2-5. habiting reef corals, III. A revision of the ---. 1980b. Coral-inhabiting crabs of the family Hapalocarcinidae (Brachyura), with family Hapalocarcinidae from Japan. III. some remarks on their morphological pecu New genus Fizesereneia. Bull. Natl. Sci. liarities. Palao Trop. BioI. Stn. Stud. 2(4): Mus. Tokyo, ser. A (Zool.) 6(3): 137-146, 687-731, figs. 1-17, pI. 3-5. figs. 1-5.