II August 2003 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON I 16(2):438-452. 2003 . Establishment of a new genus for Panopeus bermudensis Benedict & • Rathbun, 1891 and several other xanthoid crabs from the Atlantic and Pacific oceans (Crustacea: Decapoda: Xanthoidea)
Darryl L. Felder and Joel W. Martin
(DLF) Department of Biology, University of Louisiana, Lafayette, Louisiana 70504, U.S.A., e-mail: [email protected] (lWM) Natural History Museum of Los Angeles County, Los Angeles, California 90007, U.S.A., e-mail: [email protected]
Abstract.-The new genus Acantholobulus is proposed to accommodate sev eral brachyuran crab species formerly assigned to the genera Panopeus H. Milne Edwards, 1834 and Hexapanopeus Rathbun, 1898, with Panopeus ber mudensis Benedict & Rathbun, 1898 as the type species. Characters of the carapace front, anterolateral dentition, male first pleopod, and larval stages define the new genus. Morphological findings are congruent with recently re ported molecular evidence for distinction of this genus. The western Atlantic species Hexapanopeus hemphillii Benedict & Rathbun, 1891, Panopeus gatu nensis Abele & Kim , 1989 and Hexapanopeus heblingi Rodrigues & de Loyola e Silva, 1998 are considered junior synonyms of Acaruholobulus bermudensis, new combination. Hexapanopeus schmitti Rathbun, 1930 from the western At lantic is assigned to the new genus, and Panopeus margentus Williams & Boschi, 1990 is concluded to be a junior synonym of Acantholobulus schmitti, new combination. Panopeus mirafloresensis Abele & Kim, 1989 from the east ern Pacific and Panopeus pacificus Edmondson, 1931 from Hawaii and Tahiti are also assigned to Acantholobulus. While provisionally retained among the Panopeidae Ortmann. 1893. genetic studies and larval morphology distance Acantholobulus, new genus, from typical panopeid crab genera examined to date.
The genus Panopeus H. Milne Edwards, cies often facilitates ready distinction from J 834 includes a number of extremely com the larger forms, the latter of which tend to mon and abundant intertidal and upper sub share a very characteristic and conserved tidal marine and estuarine crab species, es gonopod morphology (Williams 1984£1, b) . pecially along warm-temperate to tropical A unique gonopod morphology has been re coasts of the Americas. Taxonomic history ported for Panopeus bermudensis Benedict of this group is confusing, and superficial & Rathbun, 1891 (Monod 1956). P. paci similarities in morphology of the adults of ficus Edmondson, 1931 (Forest & Guinot ten make species identifications particularly 1961) , P. gatunensis Abele & Kim, 1989 difficult. This is especially true for juveniles (Abele & Kim 1989) , P. 111 irafioresensis • of those species that attain large size as Abele & Kim, 1989 (Abele & Kim 1989), adults, since immature stages of these crabs and P. inargentus Williams & Boschi, 1990 superficially resemble adults of smaller (Williams & Boschi 1990). In addition. dif panopeid species. However. the complex ar ferences in carapace granulation and denti mature of male first pleopods (gonopods) in tion, together with the shape of the male mature individuals of the small-sized spe- abdomen. distinguish this group from most •
• VOLUME 116. NUMBER 2 439 species of Panopeus and from other pano above and below. Anterolateral teeth peid genera like Hexapanopeus Rathbun, strongly developed, prominent. arrayed in 1898, Rhithropanopeus Rathbun, 1898, Eu distinct arc, third and fourth teeth with tips rypanopeus A. Milne-Edwards, 1881, and anteriorly directed, fifth tooth well defined, Eurytium Stimpson, 1859. thick, acute. Abdomen of male with termi Earlier studies of larvae for American nus rounded or weakly triangular, "lateral panopeid crabs, which included P. bermu extremities on third segment rounded, sixth densis, revealed unique characters in zoeal segment slightly broader than long; lateral morphology for the latter species (Martin et lobe of third segment not contacting coxa al. 1984, 1985). More recently, one of us of fifth pereopod, seventh sternite distinctly (DLF) has participated in comparative ge exposed. Dactyl of major chela with strong netic studies that unambiguously position basal tooth (Fig. I b); color of immovable P. bermudensis apart from most panopeids finger extended variably onto palm; carpus for which mtl6S DNA has been sequenced distally with distinct transverse groove. (Schubart et al. 2(00). Finally, extensive Fixed finger of major and minor chela in collections from the Indian River lagoon, male deflected, less so in female. Male first Florida, the Gulf of Mexico, and several pleopods (gonopods) terminally complex Caribbean localities have facilitated studies (Fig. Ic-f', 2a-I), subterrninally with row of of adult morphology that provided insight short denticles, field or row of a few strong into variations in characters at both the spe setae, and strong subterminal tooth directed cific and generic level. In view of our pre at right angle to shaft; terminal apex (ac sent evidence from adult morphology, lar cessory process) tapered, usually acute; val characters, and molecular genetics, we soft, variously folded median lobe sur herewith establish a new genus for this spe rounding terminus of tract and bearing I to cies and its closest relati ves. several strong, distally directed spines or Materials examined include selected setae. holdings from National Museum of Natural Diagnosis of roeal morphology.-Zoeal History Smithsonian Institution, Washing development (known for the former Pano ton, D.C. (USNM), the Museum national p eus b ermudensis and Hexap ano p e us d 'Histoire naturelle, Paris (MNHN), and the schmittiy consisting of 4 stages; all stages University of Louisiana, Lafayette Zoolog Jacking lateral carapace spines or with, at ical Collections (ULLZ). Size is expressed most, a small lateral protrusion of the car as maximum carapace width (CW) mea apace where such spines exist in other xan sured in millimetcrs (mm) and includes the thoid larvae (Fig. 3a, b, e. h. i). Antennal anterolateral teeth. protopod unarmed and slightly dilated at tip; antennal exopod absent or greatly re Acantholobulus, new genus duced: antennal endopod absent or reduced (Fig. 3c, 1', j). Arms of telson furca each Type species.s--Pcmopeus bertnudensis with I dorsal spine located below (posterior Benedict & Rathbun, 1891. to) insertion point of. and sometimes pos Diagnosis oj" adult morphology.s-s-Ceie terior to, furthest distal extent of telson mar pace (Fig. Ia) moderately convex, slightly ginal setae; other telson spines may be lo elliptical in outline, regions well-defined by cated anterior to this (e.g., Fig. 3g). Other grooves, usually crossed by raised trans characters as in the "Group I" xanthid zo verse lines of granules on anterior half. eas (Rice 1980, Martin 1984. Martin et al. Frontal edge bilobed with distinct median 1985). fissure, thickened granulate margin usually EtYll1ology.-Combines prefix "acan with transverse concavity, sometimes de tho-," meaning thorny as in a thorny plant, fined as furrow between rows of granules with"lobulus," meaning small lobe, in ref- 440 PROC EEDI NGS O F T HE BIOLOGI C AL SO CIETY OF WASHINGTON
Fig. 1. AC(//1//IO!" /JII /us bcrmu dcnsis (Benedic t & Ra thbun. IW)Il. new co mbi natio n. syruyp ic spec ime ns from Bermuda. US NM 42804 . Ma le. CW 12.5 n1l11 : a. carapace and eyes. dorsal sur face ; h. maj or che la. external surface. Male. CW 7.6 mm: c-f. rig ht go no pod mesial. lateral. abdo m inal and ste rnal surfaces. respe ctively. Scale lines ind icate D.5 rnrn. erence to the median lobe of the male go n wa ters of the tropi cal ea stern Pacinc reg ion; opod in this genus, which generally bears Acan tho lo b ulus pacificus (E d mondson, characteri sti c spiniform setae . The gender is 1931 ), new combinati on, forme rly Pan o masculine. peu s paclficus, including "Neopanopc sp. Assigned sp ecies.s-Acantholobulus ber 'l " of Edmonds on, 193 J ), from Hawaii and mudensis (Benedict & Rathbun, 1891), new Tahiti ; and Acantho lobulus schmitti (Rath co mbination, formerly Pan opeus bermu bun , 1930 ), new combination , formerl y densis. se nior sy nonym of Hexapunopeus Hexapanop eus schmitti, senior sy nony m of hemphillii Benedict & Rathbun, \89\ , Pan Pan opeus margentus William s & Bosch i, op eus gatune nsis Ab ele & Kim, 1989, and \ 990, fro m warm temperate Atl antic coastal Hexapanopeus heblingi Rodrigues & de waters of South America. Loyola e Sil va ( 1998), fro m throu ghout tropi cal to warm-temperate waters of the Acu ntholobulus bermudensis (Benedic t & western Atl antic; A cantholobulus miraflo Rathbun, 1891 ), new combination resensis (Abele & Kim, 1989) , new com Figs. 1a- f, 2a-f, 3a-d bination, formerly Panopeus mirafloresen sis, including former eas tern Pacific record s Panopeus he rbstii va r. serratus Mi ers, of P. bermudensis, from throughout coastal 1886 :129. VOLUME I 16, NUMBER 2 441
Fig . 2. Acnntlioiobulus bcrnuulensis (Benedict & Rathbun. IX9 I) . new cumbination: a. b. syntypic male from Bermuda. USNM 42X04. CW 12..5 mm, right gonopod mesial and lateral sur faces. respectively (subterminal tooth broken and missing from specimen. shown as dolled line): c. male from Macau. Rio Grande do Norte. Brazil. USNM 307299. (one of several specimens reported as " Pu nop eus miraflorrsensis" by Ferreira & San karankutty 1997). CW 7..5 mm. right gonopod; d-T, male from Indian Key. Florida. US NM 1.5649 (holotype of Hcxapanopeus hell/phi/Iii Benedict & Rathbun. IXl)I). CW 9. 7 nun, right gonopod abdominal, mesial. and stern al surfaces. respectively. Acantholobulus schniitti (Rathbun. 1930), new combination: g . h. holotypc male. Rio de Janeiro. Brazil. USNM .59X31. CW 12.8 mm, right gonopod abdomina! and sternal surfa ces . respectively ; i, j . male from Escollcra Norte. Mar del Plata. Argentina. U S NM 23919 I (holotype of PI/lIOI'l'IIS nntrgentus Williams & Boschi, 1990). CW 10.6 mm, lclt gonopod sternal and abdominal surfaces. respectivel y: k, I. male from Eseollera Norte. Mar del Plata. Argentina. USNM 23919.5 (par.uype of Punopeus n/(II"'~l'II1I1S Williams & Bosehi, 1990). CW 14.X mrn , right gonopod abdominal and sternal sur faces, respectively. Scale line indicates 0 .5 mm. 442 PROCEEDINGS Or- TH E BIOLOGICAL SOCIETY OF WAS HINGTON
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Fig. 3. Zocul charac ters of Acuntho lob ulus, new ge nus. a- d . Acantholobulus bermude nsis (Benedic t & Rath bun . IH91): a. second zoeal stage. lateral view; h. fourth zoeal stage. frontal view . arrow show ing diagnostic lack of lateral ca rapace spine; c. second zoeal stage. an tenna. arrow showing diagnostic dil ated tip of protopod ; d. left tclsonal turca. dorsal view. arro w showi ng diagnostic posterior dorsal spi ne of furca l arm (from Ma rtin ct al.. 1985. as Punopeus be rmudensis Benedi ct & Rathbun . 189 \). c-g. Acuntholobulus s,hm;/li (Rathbun. 1930). all lig ures of second zoeal stage: e. front olateral view ; b. an tenna: c. telso n. dorsal view (from Bakker ct al.. 1989. as Hex upa nopeus schmitti) , h- k. A ClIlII/W /o!JU /lIs schmittl: h. seco nd zoe al stage. lateral view; i, thir d zoea l stage. fro nta l view showi ng lateral protrusion s of carapace; j . second zoeal stage. ante nna; k, second zoeal stage telson, dorsal view (from Rodri guez & Sp ivak 2(0). as Pan op cu s margcntus Williams & Boschi, 1990). NOI drawn to sca le. VOLUME /16, NUMB ER 2 443
Panopeus wurdemannii Benedict & Rath Panopeus gatunensis Abele & Kim , 1989 : bun, 1891:357,372, pl. 24, figs, 6, 7, 3. 3 1, 32, 41. fig. 15a-h, table 3. Panopeus bermudensis Benedict & Rath Panopeus permudensis (sic).-Sankaran bun, 1891 :357, 376, pl. 20, fig, 2, pl. 24, kutty & Manning, 1997 :254. figs, 14, 15,-Rathbun 1930 (part): vii. Panopeus miraftoresensis.s-s-Fevveu» & 334 (key), 360-363, 365, 394, 5lB, fig. Sankarankutty, 1997 :153-155, fig. l a-f 56, pl. 165,-Rathbun 1933:60, 62. (Brazilian specimens only: not Panopeus Monod 1956:325, 336, figs. 439-440. mirafioresensis Abele & Kim , 1989). Felder 1973:5, 69 , pI. 9, fig. 20.-Powers Hexapanopeus heblinMi.-Rodrigues & de 1977: 10 I.-Camp et al. 1977:35, 52, ta Loyola e Silva, 1998 :263-270, figs. 1 ble 36.-Gore et al. 1978:221, 224. 228. 20. 231-235, 237, 242-243, 248, table 2-4. Inot " Panope us bennudensis" of Lebour, figs. 8, 9, App, L-Felder & Chaney 1944:119, fig. 9a-dl. 1979:15.-Martin 1984 :227, 229, 230, 232 , 233 (key to zoea), table I.-Lemai Material examined.-Type series, 16 tre 1981 (part, not Paci tic range): 253 specimens, of Panopeus berniudensis from 255 , fig . 5a.-Markham & McDermott Bermuda, USNM 42804; holotype male of 1981: I273. -Martin et al. 1984 :537-542, H. hemphillii from Indian Key, Florida 544-549, 555-559, 563 -564, 577-581, (USNM 15649); holotype male of P. s« 589, 592-595, 597, 598. figs. 2-5, lOa, tun ensis from Gatun Locks of Panama Ca Ila, 12-13a, i, 17, 21-23a, i, 30-33a, i, nal, Caribbean side, Republic of Panama 43-48, tables 4. 5.-Felder et al. 1985 : (USNM 237647); 2 males, I female report 186, 208, fig. 14.-Martin et al. 1985 : ed as ..P. mirajlorcsensis" by Ferreira & 84-103, figs. 1-7, tables I , 2.-Abele & Sankarankutty (1997) from Macau , Rio Kim 1986:xv, 59, 610 (key), 630, Grande do Norte, Brazil (USNM 307299). 63Ic.-Martin & Abele 1986 : 190, fig. Diagnosis and common variations o] IK (not fig. 3C). -Martin 1988 :76 , 82, adulrs.-Carapace (Fig. I a) broadly oval; 89, 90 , fig. 2f-h, j, k, 7, table I.-Abele anterolateral teeth broadly developed, tend & Kim 1989:31, 36.-Williums & Boschi ing to lobiforrn, obtuse in small specimens, 1990:599.-Melo 1996 (part, not Pacific first and second broadly fused together with range):368 (key), 371 , 598.-Camp et al . separation sometimes obscure ; dorsal out 1998: 149, 212.-Schubart et al. 2000: line for each lobe of front arcuate near mid 1168-1169, 1171, fig. I , tables 1,3. line , becoming truncate or weakly concave Panopeus hemphillii Benedict & Rathbun, laterally. Lines of granules on carapace usu IlNI:357, 374, pl. 24, figs. 12, 13. ally including distinct short ridge (some Eupanopeus hennlldensis.-Rathbun, 1898: times paired by short parallel ridge) on ei 273.-Rathbun 190 1:29. ther half of epigastric region, composed of Hexupanopeus hemphillii Rathbun. 1898: enlarged granules about same size as those 273.-Rathbun 1901:3J.-Rathbun 1930: of hepatic or branchial ridges. Color of 384 (key), 400-402, pl. 171. figs. I, 2, movable finger (Fig. 1b) variably extended 6.-Rathbun 1933:63 .-Abele & Kim onto palm, varying from brown to tan , yel 1986:xiv. 57, 608 (key). 620. 621 c-e. lowish. or ivory; if dark. tips of fingers fad Camp et al. 1998: 148. 221. ed to near white. Palm usually with light, Eupanopeus bermudensis val'. sculptus Ver nearly white area distally near base of mov rilL 1908 :357. able finger. Gonopod (Fig. Ic-f. 2a-f) with Panopeus hermudensis'l (juvenile).-Feld median lobe of terminus variously tubi cr. 1973:62. form, slightly folded toward terminal apex Panopeus species 2.-Martin & Abele. (accessory process). not exceeding terminal 1986: 189. fig. 2B . apex distally, usually bearing 2-3 elongate, 444 PROCEED1NGS OF TH EBIOLOG ICAL SOC IET Y OF WASHI NGTON
b
Fig . 4. Typica l zoc ul charac ters of the ge nus Punopeus H. Milne Edwards, I!D4 and related genera in the family Panopcidae. a. second zoca l stage. lateral view, arrow showing we ll developed lateral carupucc spine ; h. second zoea l stage. antenna, arro w sho wing tapered distal ext remity (note presence of spines and relative exopodl endo pod sizes): c. third zoca l stage. telso n in dorsal view. arrow showing placement of furcal spi nes (redrawn from Monni et al. 199:-;. as I'lI/lOl" 'II S austrobcsns Williams 1984a ). curved spi nes. Zoeul stages (Fig . 3a- d ) o ut the Gulf of Mexi co and Caribbean Sea, with arms of telsonal Iur ca lacking lateral the Antilles, and northern coas tlines o f spination , eac h ann bearing single dorsoin South Am erica to the state of Santa Catar tcrnal spine originating posterior to distal ina . Brazil. tips of internal marginal setae of Iurca. Oc RClIlllrks.-No holotype was designated casional variants of adults include speci in the publish ed original description of men s having front a thicken ed lip, lacking Panopeus be rm ud cnsis by Benedict & transverse furrow on frontal lobes : beari ng Rathbun (18 91), though a " large male" obscurely de veloped lines of g ranules dor wa s reported be 12 mrn C Wo Subsequently, sa lly o n carapace : lacking dimorphism in Rathbun ( 1930) ind icated that the type se che lae, with both being developed as en ries consist ed of 15 specime ns "inc luding larged minor che lae lacking basal tooth on hol ot ype ' and indicated that the " type movable finger; or bearing more or less male" as 14.4 mm C Wo We presently find than 2 o r 3 spines on the median lobe of no specimen larger than a male of 12. 5 mrn the go nopod terminus. CW in the ty pe se ries o f 16 spec imens tha t D is! ributio//.-Warm-t ern pe rate and comprise USNM 428 04. We mu st sur m ise tropical sha llo w waters of the western At that the presumed ho lo type is e ith er missing lantic, from Bermuda and F lorida. through- or that the size or the largest speci men and VOLUME 116. NU MBER 2 445 presumed holotype was in error as reported consistency as to suggest tha t they are like by Rathbun ( 1930). Whi le 16 specime ns re ly characters of a separate species. main in this lot, which exceede d the num Records of Hexapanopeus hemphillii are ber reported by Rathbun, a small, possibly not to be found in the literature since Rath original tag wi th this Jot docs ap pear to in bun (1930). and ou r attempts to collect dica te that the co llec tion co nsis ts of "IS+,. from habitats described in that work con specime ns. In absence of any more defini sistently yield only specimens assignable to tive ev ide nce tha t a designated holotype Acantholobulus bermudensis. Examination ever ex isted, we regard this series as syn of the male holotype of H. he mphillii types. (USNM 15649 ) revealed tha t it is clearly Monad ( 1956 :325) first noted the po ten ass ignable to Acantho lobulus . Its gonopod tial significa nce of the go nopod of Pano (Fig. 2d. e) differs from the typical form in peus bermudensis as a ge neric- leve l cha r A. bermudensis in that it bears three smaller, ac ter. Subsequent workers also ca lled atten rath er than two larger, spi nes on the medial tion to the uni qu e term inal go nopod mo r lobe. T his co ndi tio n, however, is not un pho logy and its possib le generic co mmo n amo ng males in large sa mples of significance in bot h P. bermudensis and A. bermudensis. Also, white co loration on Panopeus mirafloresen sis (Martin &Abele the fingers of the chelipeds, re lative widths 1986; Martin et al. 1984 , 1985; Abele & of the anterolateral teeth on the carapace, Kim 1989). In addition, PWlOlJeUS gatunen and the ca rapace shape, as used in the de sis, herein recognized as a j unior sy nonym scription and later diagn osis of H. hemphil of Acantho lobulus bermudensis, was shown Iii (see Benedict & Rat hbun 189 1, Rathbun to have a go nopod simi lar to the aforeme n 1930), all can be found among larger series tio ned two species, differing primarily in of A. bermudensis. In the absence of de havi ng a supernumerary late ra l sp ine. With pendable diagnostic characters for H. hem out question, all three of these species share phillii, we must treat it as a junior synonym go nopod morphology diag nos tic of Acan of A. bermudensis. tholobulus. However, we place no signifi Likewise, no characters are offered in the ca nce in the observatio n of a supernumer description by Rodrigues & de Loyola e ary latera l spi ne on the go no pod of P. ga Silva (1998) that wo uld differentiate Hex tunensis tha t was origina lly illu strated by apanopeus heblingi from A. be rniudensis, Abele & Kim ( 1989 : fig. 15f- h) especially and inte rspecific comparisons they make since the ho lotype ma le (USNM 237647, are limited to western Atlant ic species of the only known specime n) has only a sing le Hcxupan op eus. We have not been able to lateral spine on the op pos ite go nopod fro m examine type materials of this species, but that illustrated. Furthermore, supernuma ry both the text and figures in the pu blished lateral spines are occ asionally see n among description clearly detail fea tures diagnostic large series of spe cimen s assignab le to A. of Acantho lobulus, new ge nus, we ll wi thin bermude nsis from the Indian River reg ion the range of variation tha t we have ob of Florida (UL LZ co llections); whe n these served in A. be rmudensis. At least for the occur, they are usuall y on o nly one side of present, we conclude that this species is the male. The ho lotype of P. gatunensis also a junior synonym of A. bermudensis, a also ap pea rs to have a regenerated major finding supported by the strong similar ity chela, an atypically smooth carapace, and a of its larvae (Rodrigues 1997) to those of very poorly defined transverse fro ntal fur A. bermudensis (see Discussion below). row. T hese are, however. agai n fea tures A report of Panopeus mirafioresensis see n in a small percent age of specimens as from Brazil (Ferreira & Sankarankutty signable to A. bermudensis from Florida, 1997) is inte rpreted to also represent A call and none occur in such combination and tholobulus bermudensis rather than the 446 PROCEED INGS OF T HE BIOLOG ICAL SOC IETY OF W A SH INGT ON aforementioned s pec ies , which appears to into warm-temperate waters or so uthe rn be restricted to Pacifi c coastline s , Speci Bruzil and Uruguay. lnxreud, the authors did mens upon which this Bruzifian record was note its re semblance to Panopeus bcrmu based we re furn ish ed to the late R. B . Man d CIISi.I' and the Paci fic P. miraflo rcsensis, ning prior to publication and (as not ed by both or which are in the present work es Ferreira & Sunkarunkuny 1997: 157) he tabl ished as con gen er s o r AClIllt !W!O!JIf ! IIS co uld not confir m that they represented scl uuitti. We ha ve found no consi stent fea Panopeus miruftoresen sis . We have e xam tures of ge ne ral carapace and che lipe d mor ined these specimen s (USNM 3072lJ9), il ph ology that would se rve to distinguish P. lustrated the male gonopod (F ig. 2c) and nta rg enrus from larger specime ns of Aca n found them indist ingu ishable from A. ber t hO !O /JII ! lIS schmitti. Furthermore. the right mudcnsis. We also observe, howe ver, that gon o pod as figured in the origina l descrip the Paci lie speci es Acantholobulus ntirajlo tion (Williams & Boschi 1990: fig. l e, d ) resensis (as herein assi gned) is itself not appears to be highly aberrant and perhaps readily distinguishabl e from A. bermudcnsis damaged . Our own e xa mi natio n of the on the basis of cha rac te rs reported to date, damaged holotype rev ealed the left gono es pecia lly given the ran ge of variatio n that pod o f th is specime n to be intac t (Fig. 2i . we have observed in populations of the lat j) an d almost identic al in term inal mor ter spe cies. Records from the eastern Pacifi c ph ol ogy to that herein figure d fo r the ho regi on inc lude mention of both Panopeu s lotype of A{,Ollt ho!OIJII ! llS schntitti (Fig, 2g, be rnuulens is (see Rathbun 1930, Garth h). Howev er, as discuss ed bel ow. the de 1961. Lc maitre 199 I. Abele & K irn 19X9, scription of larval development for thi s spe Lemaitre & Alvarez Leon 1( 92) and P. cies ( P o II O!JeIlS niargcntus) hy Rodrfguez & niiraflorcscnsis (see A be le & Kim 19W): Spivak (200 1) included some differences 36), and no atte mpt is herein mad e hy us to from tha t presented by Bakke r et <.II. ( 1l)X9) resolve morphologically wheth er these east for H cxap unopeu s schmitti, w hich sho uld e r n Pu c ific populations could represent not be the case if these tw o nam es ind eed more th an one spe cies. At least until further refer to the same species (fur the r addressed comparative study is undertaken and on in Discussion below). going mol ecular analyses are completed , Materials o f Panopeu s [JOCi/jCllS from we recognize A. mirafiorescnsis as the eas t Ha waii and Tahiti. alo ng w ith so me qu es ern Pac ific cognate o f A. bertnuden sis. The tion a bly assigne d to "Ncopauope sp. T' by natural ran ge of latt er species appears lim Edmondson «(93 1), we re re viewed by For ited to warm-temperate a nd tropical sha llo w es t & G uinot ( 1'16 1) who provided detailed waters of the western A tlantic. as also con illu strations of gonopods , chelae, and ca ra cluded by Salgado Barragan & Hendrickx pace dentition for the se spec imens, all or ( 1997). wh ich appear to be typical of Aca ntliolob Systematic review of congcl/e rs.- Re<: U!IIS. Our di rect ex amination o r topotypic ognition of Panopcus murgentus as a junio r specimens of P. pacificus (four ma les , two synonym of A cantholo bulus schntitti (Ra th fema les ) collec ted by C. H. Ed monds on bun, 1930 ), new combination. was based from Pearl Harbor (U SNM l)6304) contirrns up on exam ina tion of th e holotype (USNM this generic rea ssignment. We hav e also re 239191) and paratypes (USNM 239195) o f examined the two males o f P. pucificus re POI lOp eus m argentus, all from Mar del Pla ported from Tahiti (MNHNB 139(3) by ta. A rgentina. The description o r P. mllr For est & Guinot ( 196 1) and found no ch ar gcntus by Will iams & Boschi ( 1990) in acters tu distingu ish them from the Ha wa i eluded no co m pa riso ns to or mention of ian ma ter ia ls. Ed mondson ( 1931) preceded Hexapanopeus sc hmitti, even though the his ori ginal description or P. pacificus by latter species has been documented to range noting the po ssibility of this species being VOLUME 116. NUMBER 2 447
"transported to Hawaii through shipments Panopeus mirafioresensis, Abele & Kim, of oysters or on the bottoms of ships," es 1989: fig 18) from Panama. pecially since he found it only in associa Our proposed revisions are summarized tion with "sponges, barnacles and tunicates as follow: attached to buoys and floats in Pearl Harbor, Former name Oahu." This is typical habitat for most of New name the known Acantholobulus spp., and may Panopeus bermudensis indicate a high potential for their dispersal Acantholobulus berniudensis. and introduction among fouling materials. new combination At very least, it could account for wide dis Hexapanopeus schmitti tributions, such as that for A. pacificus, Acantholobulus schmitti, which appears to encompass both Hawaii new combination and Tahiti. However, except for the appar Panopeus mirajloresensis ent larger size evident in most specimens of Acantholobulus mirafloresensis, A. paclficus, there are also no obvious mor new combination phological characters to distinguish it from Panopeus niargentus A. mirafioresensis. More definitive conclu junior synonym of A. schmitti sions about the relationship between these Panopeus gatunensis species and their relationships to others of junior synonym of A. bermudensis the genus should be facilitated by molecular Hexapanopeus hemphiIIii genetic studies that we have in progress. junior synonym of A. bcrmudensis While we have not re-examined materials Hexapanopeus heblingi of the "Neopanope sp. '1" reported by Ed junior synonym of A. bermudensis mondson (1931: 14), from Pearl Harbor, the chelae of one large male in the lot of Pan Discussion opeus /)[IC(/;CU.\· that we examined (USNM The use of brachyuran larval morpholo 963(4) appears to match Edmondson's de gy, especially characters of the first zoeal scription of the chelae in his specimen of stage, in elucidating systematic and phylo "Neopanope sp. '1". In addition, the figure genetic questions is now relatively well es of the gonopod for"Neopanope sp. '1" pro tablished (e .g., see references in Clark et 'II. vided by Forest & Guinot (1961: fig. 1(6) 1999) . Larval morphology has been of par is clearly typical of Acantholobulus and ticular value for resolving relationships well within the range of variation to be ex within the enormous superfa m ily Xanthoi pected in gonopod morphology for species dea Macl.eay, 183R (e .g., Clark & Al-Ai in this genus. We thus conclude that the pe daroos 1996, Clark & Galil 1998, Clark & culiar cheliped dentition reported for "Neo Ng 1998, Fransozo et al. 200 l , and papers panope sp .?" by Edmondson (1931) is like cited therein). Several workers (Rice 1980; ly the result of cheliped regeneration and Martin 1984, 1988), have proposed recog that these materials represent additional nizable groupings of xanthoid larvae in at specimens of A. pacificus, new combina tempts to address systematic or phyloge tion. Cheliped variations like those ob netic questions. Within the xanthoid family served by Edmondson me not uncommon Panopeidae Ortmann, 1893 sensu Guinot in populations of A. bermudensis that we 1978 (see also Martin & Davis 200 I), lar have sampled in Florida (ULLZ collec val morphology is diverse. Prior to 1985, tions). These have also been illusrrated for however, all known species of the Panopei variant specimens of both A. bertnudensis dae save Panopeus bermudensis were (as Panopeus gatunensis, Abele & Kim, thought to have lateral carapace spines, 1989: fig. 15) and A. mirafioresensis, (as which is true of nearly all known xanthoid 44li PROCEEDI NGS OF THE BIOLOGICAL SOCI ETY OF W/\S H INGTON
lar vae . " Ty pica l" zoeal larvae of pan op eid Panop eus margcntus with Acantholobulus crabs ha ve we ll de vel oped (a ltho ug h so me schmitti , and co nc lus io n th at Hcxapan op cus times sho rt) lateral carap ac e spi nes. a ta licmphillii and H Iicbling! mu st be sy no n pering a nte nnal pro topod th at is o fte n ym ize d w ith Acam holo bulus bonnuden sis. arm ed di stally (and is never distally inffut I n the case o f Pan opcu s margentus, o ur rea ed), and tw o or three sma ll spines on the so ns for placi ng th is species into sy no ny my tel son al furcae, all located at approxima te ly wi th Acant ho lobulns sclunitti on the basis the le vel o f inse rtion of the three paired tel of adult features have been given earlier. sona l se tae (Fig. 4 ). Th e larvae o f P. ber O ne wo uld thu s expect tha t re ported lar val II I/U/CIl S ;S were described as differing from descriptions would prove to be identical, all other pan op eid lar vae in lack ing these a nd to large extent this is true: Rodrigue z la teral ca rapace spines (Martin e t al. 19R5), & Spivak (200 I :XI X) state that " Pau opeus having an ante nna that is d ilated at the tip m arg en/lis larva e are very simila r to those and that lack s a n exopocl (ac tually present, of H. sclt ntitti." T he lar vae of P. rna rgentus but greatly reduced, see Discussion), and in as desc rib ed by Rodrigu ez & S piva k (2 ()() I ) bearin g a sin gle pair of tel son al spines lo also lack lateral carapace spines. altho ug h a ca ted posteri or to the distal tip o f the inte slight protrus ion or the ca rapace ca n be rior telson al setae. Remark abl y, the zoeal seen (F ig. I h. i), w hic h may be a homo stages of Hexap un opeus schmitti as de logue of the lateral spine in typical pano sc ribed by Bakker e t al. ( 1989) also lack a peid zoeas. As de scribed, however. the lateral ca ra pace spine, have an ante nna ! zoeal sta ges of P. margen/us would appear protopod tha t is dilated at the tip, have a to lack the second pair of telson furcal red uced anten na l cxopod, a nd bear a pair of spi nes shown by Bakker e t al. ( 19X9) for posterior spi nes on the tel son al Iurca (a l H. schmitri. though. in addition. there is also a pai r o f In the course of our studies , we ha ve re more ant eri or furcal spines). exami ne d th e parental femal e (US NM O ur decisi on to group the for me r P OIlO 2R~Hl46), as well as the lar val stages of P. p eus bermudcnsis a nd H exap unopcus ma rg entus (USNM 291 175) described by schniitti. along wi th the former Pan opeus Rodrtgucz & Spivak (200 I). We ca nno t de mirafio rescn sis and P. pacificus. int o A ClIJI finitive ly separate the female from those of th olobulus. new gc nus, on the basis o f adult A. srhmitti, though females of both of these morphol ogy is thu s supporte d by lar val species re mai n poorly described. Among morph ology for at least the first two of the larvae, w he n mounted in glycerin a nd these species . We ex pec t th at lar val cha r ex amined with differential interference con ac ters of the former P. m irafioresensis and trast (DIC) optics , we do find zoeas at stag P. pac ificus, which remain und escribed , es on e throu gh four in whi ch a sec ond. ve ry sho uld resemble those of th e tw o other spe sma ll pair of telson furca l spines can be dis c ies of Acuntholo b ulus. Given the similar cerned at high magnific ati on s (first ca lle d to ities in ad ult morph ology herewith reported our atte ntio n by J. C uesta and B. Maho n) . fo r pr op osed members of Acunthoiobul us. These. however. appear to be less conspic new ge nus. shared characters in larval mor uou s than those illustrated for zoeal stages phology to the ex te nt kn own within thi s of A . schmitti by Bak ke r et al. ( f 9RlJ). and gro up, and the es tab lis he d genetic dist inc it should be no ted tha t they were not ob tio n for at least one of its members (Schu vio us on all specime ns exam ined ; in some bart et al. 2000), the ge ne ric revision herein cases, the sm all se ta ) sha ft appeared ins tead pro posed is strongly supported. as a clus ter of two or three miniscule setae Even so. so me problems remain in our or setal fragments and may ha ve been worn ass ig nme nt of species to the new ge nus or dam aged . A cantholobulus, deci sion to synonymize Fo r Hcxapanopeus he bling ! we ex am ined VOLUME I 16. NUMBER 2 449 selected figures of the larvae illustrated by quences and larval morphology set this Rodrigues (1997, figs. 3-10). On the basis group apart from typical panopeid species, of these larval descriptions, zoeal stages of and definitive assignment must await im H. heblingi are essentially identical to those proved understanding of familial separa of Panopeus be rm udensis described by tions among the Xanthoidea. Martin et al. (1984, (985). The absence of a lateral carapace spine, the inflated tip of Acknowledgments the antenna, the reduced antennal exopod, We gratefully acknowledge the advice and other features are a nearly perfect and encouragement of the late R. B. Man match, supporting our synonymy of the two ning, who assisted us in early phases of this species under the name Acantholobulus project. We also thank K. Anger, J. Cuesta, bermudensis. Yet, descriptions of the me D. Guinot, R. Lernaitre, B. Mahon, F. Man galopa appear to differ slightly. In the me telatto, K. Reed, and C. Schubart, who in galopa of P. bermudensis, described by various ways facilitated our completion of Martin et al. (1984, 1985), the frontal re this paper. Support was provided from Na gion of the carapace bears anterolateral tional Science Foundation grants no . DEB horns, and the cheliped bears a distinct re 9972100 and DEB 9978193, U.S. Depart curved spine on the ischium. Neither of ment of Energy grant no . DE-FG02 these characters is illustrated by Rodrigues 97ER 12220, ancl several small grants from (1997) for the species therein referred to as the Smithsonian Marine Station at Ft. H. heblingi. Although frontal horns on the Pierce, Florida. This is contribution No. 94 carapace are easily missed, especially if the of the UL Lafayette Laboratory for Crus carapace front is strongly deflected down tacean Research and contribution No. 547 ward, the cheliped spine is a feature that of the Smithsonian Marine Station at Ft. rarely escapes notice. Pierce, Florida. We cannot further comment on this ap parent discrepancy without directly re-ex Literature Cited amining the larvae referred to as H. heblin gi. Minute structures are easily overlooked, Abele. L. G oo & W. Kim. 1986. An illustrated guide to and it is pertinent to note that Martin et al. the marine decapod crustaceans of Floridu. (1985) also overlooked a diagnostic struc State of Florida Department of Environme ntal Regulation Technical Series . Vol. R. no . I (in 2 ture in the course of describing larvae that parts) . 760 p. we now refer to Acantholobulus bermuden Abele. L. G oo & W. Kim . 19R9. The decapod crusta sis . We have re-examined some of the ac ceans of the Panama Canal.-Smithsonian Con tual larvae treated in that earlier study tributions to Zoology 4R2 : I-50. (ULLZ, uncatalogued slide mounts), and Bakker. C. de, M. Morini. K. Anger. & L. L. Fernan des. 19R9. Larval development of Hexapano we must now report that the antenna does peus s('III11;1I; Rathbun. 19JO (Decapoda, Brach in fact possess a small exopod. It is entirely yura. Xaruhidae) reared in the laboratory. possible that other small features have also Ncrftica 4:U7-IM. escaped the notice of previous workers, and Benedict. J. Eoo & M. J. Rathhun. IR91. The genus a thorough reinvestigation of larval mor pl/I/(}pells.-Proeeedings of the U.S . Natrona! Mu seum 14:J55-JR5. plates 19-24. phology in all of these xanthoid groups is Camp. D. x.. N. H. Whiting. & R. E. Marrin. 1977. warranted. Nearshore marine ecology at Hutchinson Island. For the present, pending additional mo Florida: 1971-1974. V. Arthropods.c-sf'lorida lecular studies to build on those of Schubart Marine Research Publications. Department of et al. (2000). we continue to treat species Natural Resources 25:i . 1-6J. Camp. D. x ., W. G . Lyons. & T. H. Perkins. 199R. herewith assigned to A cantholobulus, new Checklists of selected shallow-water marine in genus, as members of the Panopeidae. As vertebrates of Florida.-Florida Marine Re noted above. however, both mtDNA se- search Institute Technical Report TR-J. Florida 4S0 PROCEEDINGS OF THE BIOLOGICAL SOCIETY O F WASH INGTON
Depa rtment o f E nviro nmental Protectio n, 1998: first zocul slage o f Platvpodirll« spcct ubilis i-xv. 1- 238. (He rbst. 1794 ) (Dccapuda . Brach yu ru. X umhi Clark. P. F.. & 1\ . M . A l-Aidaroos. 19l)fl. T he firs t /0 dac) o btained in the lah o rarory.- Gulf an d Ca cas o f Actucodrs hirs utissimus ( Ruppe!l. I X.,O) ribbcun Research 1.I:79- R5. and A. /(J 1IIXanthidae: Fran sozo. I<)t)0. I.ar va l development of l lrxa A ctacinac j.c--dou ruul of King A bd u luziz Uni I JiIl/ OPt'I/.'· puulen» [s Rathbun, 19.10 (Crustuccn, versi ty Marine Sciences 7, S pec ia l Issue : Sym Bruchy ura . X a nrhidac) under laho rato ry condi posiurn on the Red Sea Marin e Env iro nment: tio ns.- Rev ista Brusi lcira i'.oo lugia 7:31 - 4S. 20 7-2 14. Garth. J. S . 1961 . P,ln 2: Brac hygnurha Brachy rhy n Clark. I ~ J ~ . D. K . Cn luz. ms, & G . W. Po hle. 191JX. cha o III Ea stern Paci fic expeditions or the Ne w Accuracy and stu ndardiza rio n of brachy ura n la r York Zoolog ical So c iety. XI .V: Non-inter tida l val desnip tions .-Invertebrate Reproduc tion brachygnarhous c ra bs fro m the west Coast of and Development :13: 127 - 144 . Tropical A me rica.- i'.oo log iea (N ew York ) 46 : Clark. P. E . & B. S . Galil. 1')9 8. The firs t stag e zo ea '-' .1-1 S'). I plate . of Psrtulo tirnncra s!, el"iosl/ (Da'1
vae of xa nthid crabs. with a key to the known L. Fernandes. 1988. Larval devel opme nt of the xunthid zoeas of the western Atlantic a nd Gulf Brazili an mud crab P,1II0pellS aus trobesus Wil of Me xico.-Bulletin of Marin e Sci en ce 34: liams. 1983 (D ecupodu: Xanthida e) reared in 220-23 9. the laborator y.-Jou rnal of Crus tacean Biol ogy Martin. J. W. 1988. Ph ylogen eti c significance of the 8:594-613. bra chyuran me gal op a: ev ide nce from the Xan Ortmann . A. 1893. Aht he ilung : Brach yura ( Brac hy ura th idae.-Symposia of the Zoologi cal Society of gc nuina Boas). II. Unterabrhcil ung: Ca ncro idea, London 59:69-102. 2. Secti on : Ca nc rinea , I. Gruppe: Cyclom etopa. Marti n. J. W.. & L. G . Abele. 19R6. Notes on the mal e Die Dccap od en-Krebse des Surussburger Mu ple opod morphology in the bra ch yuran c rab scums. mit beson derer Berucksichtigun g del' family Panopeidae Ortmann. 189 3. sensu Guin vo n Herrn Dr. Dod erlcin hei Jap an und bei den ot (1978) (Decapoda) .-Crus taceana 50 : 182 Liu- Kiu-Inseln gcsam melte n und zur Ze it im 198. Strass burge r Mu seum aufbewahrten Fo rmcn, Martin. J. W.. & G. E. Davi s. 200 1. An updated clas VII. Theil.- Zoologische Jarbtichcr, Abthe ilung sifica tion of the Recent Crustacea-c-Science Se fur Sy stem atik , Geogra phie und Biologic del' ries 39. Natural History Museum of Los An Thiere 7:411 -495, piare 17. geles County. Powers. L. 1977. A catalogue and bibliography to the Martin. J. w.. D. L. Felder. & F. M. Truesdale. 1984 . crabs (Bruchyura) of the Gulf of Mexico. A com parative study of morphol og y and ontog Contributions in Marine Science 20(supple eny in juvenile stages of four west ern Atlantic ment):1-190. xanthoid crabs (C rustacea: Dec ap oda : Brach Rathbun. M . J. 1898. The Bruchyura of the biol ogi cal yu ra).-Philosophical Tran sactions of the Royal expedition to the Florida Key s and the Bahamas So ciety of London . se ries B. 30 3:537- 604. in 189J .-Bul/et in from the Laborntorics of Martin. J. W.. F. M. Truesda le. & D. L. Feld er. 1985. Natural History of the Slate Unive rsity of Iow a Lar val de vel opment of Pun opeu s bennude nsis 4(3):250 - 294. plates 1- 9. Benedict and Rathbun. 1891 (Brac hyu ra, Xan Rathbun. M . J. 190 1. The Br ach yura and Macrura of thid ae) with notes on zoeal cha rac te rs in xanthid Port o Rico.-BulJetin of the Uni ted Sta tes Fis h crabs.- Jo urna l of C rus tacean Biology 5:84 Co mmission for 190020(2):1-1 37. 105. Rathbun. M. J. 1930. T he ca nc roid cra bs of America Mcl.euy, W. S. 1838. On the Bachyurou s decapod of the fami lies Eurya lidae. Portunidae. Atcle Crustacea brough t from the Cape by Dr. Smith cyc lidae, Ca ncri dae, and Xanl hida e.-Bul/et in [il/ell/ded within Illu stration s of the Annulosa of of the U.S. Nationa l Mu seum 152: 1- 609. So uth Africa]. Pp . 53-72. plate 3 in A. Smith. Rathbun. M. J. 1933. Brachyuran crabs of Port o Rico vol. V. Hlu strutions of th e Zoology o f So uth A I ~ and the Virg in Island s. III Scien tific Survey of rica. In vcrtebratac . POl1o Rico an d the Virgi n Island s.- Nc w York Melo, G . A. S. 1998. Malacostr aca-s-Euca rida. Brach Aca de my of Sc iences 15(1):1- 121. yura. Ox yrh yncha and Brach yrhyn cha . Pp. 455 Rice. A. L. I
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