11 August 2003 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 116(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] (JWM) 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 hemphilUi Benedict & Rathbun, 1891, Panopeus gatu- nensis Abele & Kim, 1989 and Hexapanopeus heblingi Rodrigues &. de Loyola e Silva, 1998 are considered junior synonyms of Acantholobulus 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 miraftoresensis 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 1834 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 1984a, 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. miraftoresensis of those species that attain large size as Abele & Kim, 1989 (Abele & Kim 1989), adults, since immature stages of these crabs and P. margentus 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. lb); 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. 2000). 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-l), subterminally 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 1 to cies and its closest relatives. several strong, distally directed spines or setae. Materials examined include selected holdings from National Museum of Natural Diagnosis of zoeal morphology.—Zoeal History Smithsonian Institution, Washing development (known for the former Pano ton, D.C. (USNM), the Museum national peus bermudensis and Hexapanopeus d'Histoire naturelle, Paris (MNHN), and the schmitti) consisting of 4 stages; all stages University of Louisiana, Lafayette Zoolog lacking 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 millimeters (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 duced; antennal endopod absent or reduced Acantholobulus, new genus (Fig. 3c, f, j). Arms of telson furca each Type species.—Panopeus bermudensis with 1 dorsal spine located below (posterior Benedict & Rathbun, 1891. to) insertion point of, and sometimes pos Diagnosis of adult morphology.—Cara• terior to, furthest distal extent of telson mar pace (Fig. la) 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 1" 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 Etymology.—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 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
C-f Fig. 1. Acantholobulus bermudensis (Benedict & Rathbun, 1891), new combination, syntypic specimens from Bermuda, USNM 42804. Male, CW 12.5 mm: a, carapace and eyes, dorsal surface; b, major chela, external surface. Male, CW 7.6 mm: c-f, right gonopod mesial, lateral, abdominal and sternal surfaces, respectively. Scale lines indicate 0.5 mm. erence to the median lobe of the male gon waters of the tropical eastern Pacific region; opod in this genus, which generally bears Acantholobulus pacificus (Edmondson, characteristic spiniform setae. The gender is 1931), new combination, formerly Pano masculine. peus pacificus, including ''Neopanope sp. Assigned species.—Acantholobulus ber ?" of Edmondson, 1931), from Hawaii and mudensis (Benedict & Rathbun, 1891), new Tahiti; and Acantholobulus schmitti (Rath combination, formerly Panopeus bermu bun, 1930), new combination, formerly densis, senior synonym of Hexapanopeus Hexapanopeus schmitti, senior synonym of hemphillii Benedict & Rathbun, 1891, Pan Panopeus margentus Williams & Boschi, opeus gatunensis Abele & Kim, 1989, and 1990, from warm temperate Atlantic coastal Hexapanopeus heblingi Rodrigues & de waters of South America. Loyola e Silva (1998), from throughout tropical to warm-temperate waters of the Acantholobulus bermudensis (Benedict & western Atlantic; Acantholobulus miraflo- Rathbun, 1891), new combination resensis (Abele & Kim, 1989), new com Figs, la-f, 2a-f, 3a-d bination, formerly Panopeus mirafloresen- sis, including former eastern Pacific records Panopeus herbstii var. serratus Miers, of P. bermudensis, from throughout coastal 1886:129. VOLUME 116, NUMBER 2 441
Fig. 2. Acantholobulus bermudensis (Benedict & Rathbun, 1891), new combination: a, b, syntypic male from Bermuda, USNM 42804, CW 12.5 mm, right gonopod mesial and lateral surfaces, respectively (subterminal tooth broken and missing from specimen, shown as dotted line); c, male from Macau, Rio Grande do Norte, Brazil, USNM 307299, (one of several specimens reported as "Panopeus mirafloresensis" by Ferreira & San- karankutty 1997), CW 7.5 mm, right gonopod; d-f, male from Indian Key, Florida, USNM 15649 (holotype of Hexapanopeus hemphillii Benedict & Rathbun, 1891), CW 9.7 mm, right gonopod abdominal, mesial, and sternal surfaces, respectively. Acantholobulus schmitti (Rathbun, 1930), new combination: g, h, holotype male, Rio de Janeiro, Brazil, USNM 59831, CW 12.8 mm, right gonopod abdominal and sternal surfaces, respectively; i, j, male from Escollera Norte, Mar del Plata, Argentina, USNM 239191 (holotype of Panopeus margentus Williams & Boschi, 1990), CW 10.6 mm, left gonopod sternal and abdominal surfaces, respectively; k, 1, male from Escollera Norte, Mar del Plata, Argentina, USNM 239195 (paratype of Panopeus margentus Williams & Boschi, 1990), CW 14.8 mm, right gonopod abdominal and sternal surfaces, respectively. Scale line indicates 0.5 mm. 442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Zoeal characters of Acantholobulus, new genus, a-d, Acantholobulus bermudensls (Benedict & Rath- bun, 1891): a, second zoeal stage, lateral view; b, fourth zoeal stage, frontal view, arrow showing diagnostic lack of lateral carapace spine; c, second zoeal stage, antenna, arrow showing diagnostic dilated tip of protopod; d, left telsonal furca, dorsal view, arrow showing diagnostic posterior dorsal spine of furcal arm (from Martin et al., 1985, as Panopeus bermudensls Benedict & Rathbun, 1891). e-g, Acantholobulus schmltti (Rathbun, 1930), all figures of second zoeal stage: e, frontolateral view; b, antenna; c, telson, dorsal view (from Bakker et al., 1989, as Hexapanopeus schmltti). h—k, Acantholobulus schmitti: h, second zoeal stage, lateral view; i, third zoeal stage, frontal view showing lateral protrusions of carapace; j, second zoeal stage, antenna; k, second zoeal stage telson, dorsal view (from Rodriguez & Spivak 2001, as Panopeus margentus Williams & Boschi, 1990). Not drawn to scale. VOLUME 116, NUMBER 2 443
Panopeus wurdemannii Benedict & Rath- Panopeus gatunensis Abele & Kim, 1989: bun, 1891:357, 372, pi. 24, figs. 6, 7. 3, 31, 32, 41, fig. 15a-h, table 3. Panopeus bermudensis Benedict & Rath- Panopeus permudensis (sic).—Sankaran- bun, 1891:357, 376. pi. 20. fig. 2. pi. 24, kutty & Manning, 1997:254. figs. 14, 15.—Rathbun 1930 (part): vii, Panopeus mirafloresensis.—Ferreira & 334 (key), 360-363, 365, 394, 583, fig. Sankarankutty, 1997:153-155, fig. la-f 56, pi. 165.—Rathbun 1933:60, 62.-- (Brazilian specimens only; not Panopeus Monod 1956:325, 336, figs. 439-440.— mirafloresensis Abele & Kim, 1989). Felder 1973:5, 69, pi. 9, fig. 20.^Powers Hexapanopeus heblingi.—Rodrigues & de 1977:101.—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, [not "Panopeus bermudensis" of Lebour, figs. 8, 9, App. I.—Felder & Chaney 1944:119, fig. 9a-dl. 1979:15.—Martin 1984:227, 229, 230, 232, 233 (key to zoea), table 1.—Lemai- Material examined.—Type series, 16 tre 1981 (part, not Pacific range): 253- specimens, of Panopeus bermudensis from 255, fig. 5a.—Markham & McDermott Bermuda, USNM 42804; holotype male of 1981:1273.—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. ga 589, 592-595, 597, 598, figs. 2-5. 10a, tunensis from Gatun Locks of Panama Ca Ua, 12-i3a, i, 17, 21-23a, i, 30-33a, i, nal, Caribbean side, Repubfic of Panama 43-48, tables 4, 5.—Felder et al. 1985: (USNM 237647); 2 males, 1 female report 186, 208, fig. 14.—Martin et al. 1985: ed as "P. mirafloresensis" by Ferreira & 84-103, figs. 1-7, tables 1, 2.—Abele & Sankarankutty (1997) from Macau, Rio Kim 1986:xv, 59, 610 (key), 630, Grande do Norte, Brazil (USNM 307299). 631c.—Martin & Abele 1986:190, fig. Diagnosis and common variations of IK (not fig. 3C).—Martin 1988:76, 82, adults.—Carapace (Fig. la) broadly oval; 89, 90, fig. 2f-h, j. k, 7, table 1.—Abele anterolateral teeth broadly developed, tend & Kim 1989:31, 36.—Williams & Boschi ing to lobiform, 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. 1, tables 1, 3. line, becoming truncate or weakly concave Panopeus hemphillii Benedict & Rathbun, laterally. Lines of granules on carapace usu 1891:357, 374, pi. 24, figs. 12, 13. ally including distinct short ridge (some Eupanopeus hemiudensis.—Rathbun, 1898: times paired by short parallel ridge) on ei 273.—Rathbun 1901:29. ther half of epigastric region, composed of Hexapanopeus hemphillii Rathbun, 1898; enlarged granules about same size as those 273.—Rathbun 1901:31.—Rathbun 1930; of hepatic or branchial ridges. Color of 384 (key), 400-402, pi. 171. figs. 1, 2, movable finger (Fig. lb) variably extended 6.—Rathbun 1933:63.—Abele & Kim onto palm, varying from brown to tan, yel 1986:xiv, 57, 608 (key), 620, 621c-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 var. sculptus Ver- nearly white area distally near base of mov rill, 1908:357. able finger. Gonopod (Fig. Ic-f, 2a-f) with Panopeus bermudensis! (juvenile).—Feld median lobe of terminus variously tubi- er, 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 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Typical zoeal characters of the genus Panopeus H. Milne Edwards, 1834 and related genera in the family Panopeidae. a, second zoeal stage, lateral view, arrow showing well developed lateral carapace spine; b, second zoeal stage, antenna, arrow showing tapered distal extremity (note presence of spines and relative exopod/ endopod sizes); c, third zoeal stage, telson in dorsal view, arrow showing placement of furcal spines (redrawn from Montii et al. 1998, as Panopeus austrohesus Williams 1984a). curved spines. Zoeal stages (Fig. 3a-d) out the Gulf of Mexico and Caribbean Sea, with arms of telsonal furca lacking lateral the Antilles, and northern coastlines of spination, each arm bearing single dorsoin- South America to the state of Santa Catar- ternal spine originating posterior to distal ina, Brazil. tips of internal marginal setae of furca. Oc Remarks.—No holotype was designated casional variants of adults include speci in the published original description of mens having front a thickened lip, lacking Panopeus bermudensis by Benedict & transverse furrow on frontal lobes; bearing Rathbun (1891), though a "large male" obscurely developed lines of granules dor- was reported be 12 mm CW. Subsequently, sally on carapace; lacking dimorphism in Rathbun (1930) indicated that the type se chelae, with both being developed as en ries consisted of 15 specimens "including larged minor chelae lacking basal tooth on holotype" and indicated that the "type movable finger; or bearing more or less male" as 14.4 mm CW. We presently find than 2 or 3 spines on the median lobe of no specimen larger than a male of 12.5 mm the gonopod terminus. CW in the type series of 16 specimens that Distribution.—Warm-temperate and comprise USNM 42804. We must surmise tropical shallow waters of the western At that the presumed holotype is either missing lantic, from Bermuda and Florida, through or that the size of the largest specimen and VOLUME 116, NUMBER 2 445 presumed holotype was in error as reported consistency as to suggest that they are like by Rathbun (1930). While 16 specimens re ly characters of a separate species. main in this lot, which exceeded the num Records of Hexapanopeus hemp hill ii are ber reported by Rathbun, a small, possibly not to be found in the literature since Rath original tag with this lot does appear to in bun (1930), and our attempts to collect dicate that the collection consists of "15 + " from habitats described in that work con specimens. In absence of any more defini sistently yield only specimens assignable to tive evidence that a designated holotype Acantholohulus hermudensis. Examination ever existed, we regard this series as syn- of the male holotype of H. hemphillii types. (USNM 15649) revealed that it is clearly Monod (1956:325) first noted the poten assignable to Acantholohulus. Its gonopod tial significance of the gonopod of Pano- (Fig. 2d, e) differs from the typical form in peus hermudensis as a generic-level char A. bermudensis in that it bears three smaller, acter. Subsequent workers also called atten rather than two larger, spines on the medial tion to the unique terminal gonopod mor lobe. This condition, however, is not un phology and its possible generic common among males in large samples of significance in both P. hermudensis and A. hermudensis. Also, white coloration on Panopeus mirafloresensis (Martin & Abele the fingers of the chelipeds, relative widths 1986; Martin et al. 1984, 1985; Abele & of the anterolateral teeth on the carapace, Kim 1989). In addition, Panopeus gatunen- and the carapace shape, as used in the de sis, herein recognized as a junior synonym scription and later diagnosis of H. hemphil- of Acantholohulus bermudensis, was shown Hi (see Benedict & Rathbun 1891, Rathbun to have a gonopod similar to the aforemen 1930), all can be found among larger series tioned two species, differing primarily in of A. hermudensis. In the absence of de having a supernumerary lateral spine. With pendable diagnostic characters for H. hem out question, all three of these species share phillii, we must treat it as a junior synonym gonopod morphology diagnostic of Acan of ^. hermudensis. tholohulus. However, we place no signifi Likewise, no characters are offered in the cance in the observation of a supernumer description by Rodrigues & de Loyola e ary lateral spine on the gonopod of P. ga- Silva (1998) that would differentiate Hex tunensis that was originally illustrated by apanopeus hehlingi from A. bermudensis, Abele & Kim (1989: fig. 15f-h) especially and interspecific comparisons they make since the holotype male (USNM 237647, are limited to western Atlantic species of the only known specimen) has only a single Hexapanopeus. We have not been able to lateral spine on the opposite gonopod from examine type materials of this species, but that illustrated. Furthermore, supernumary both the text and figures in the published lateral spines are occasionally seen among description clearly detail features diagnostic large series of specimens assignable to A. of Acantholohulus, new genus, well within bermudensis from the Indian River region the range of variation that we have ob of Florida (ULLZ collections); when these served in A. bermudensis. At least for the occur, they are usually on only one side of present, we conclude that this species is the male. The holotype of P. gatunensis also a junior synonym of A. bermudensis, a also appears to have a regenerated major finding supported by the strong similarity chela, an atypically smooth carapace, and a of its larvae (Rodrigues 1997) to those of very poorly defined transverse frontal fur A. bermudensis (see Discussion below). row. These are, however, again features seen in a small percentage of specimens as A report of Panopeus mirafloresensis signable to A. hermudensis from Florida, from Brazil (Ferreira & Sankarankutty and none occur in such combination and 1997) is interpreted to also represent Acan tholohulus hermudensis rather than the 446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
aforementioned species, which appears to into warm-temperate waters of southern be restricted to Pacific coastlines. Speci Brazil and Uruguay. Instead, the authors did mens upon which this Brazilian record was note its resemblance to Panopeus bermu based were furnished to the late R. B. Man densis and the Pacific P. mirafloresensis, ning prior to publication and (as noted by both of which are in the present work es Ferreira & Sankarankutty 1997:157) he tablished as congeners of Acantholobulus could not confirm that they represented schmitti. We have found no consistent fea Panopeus mirafloresensis. We have exam tures of general carapace and cheliped mor ined these specimens (USNM 307299), il phology that would serve to distinguish P. lustrated the male gonopod (Fig. 2c) and margentus from larger specimens of Acan found them indistinguishable from A. ber- tholobulus schmitti. Furthermore, the right mudensis. We also observe, however, that gonopod as figured in the original descrip the Pacific species Acantholobulus miraflo tion (Williams & Boschi 1990: fig. le, d) resensis (as herein assigned) is itself not appears to be highly aberrant and perhaps readily distinguishable from A. bermudensis damaged. Our own examination of the on the basis of characters reported to date, damaged holotype revealed the left gono especially given the range of variation that pod of this specimen to be intact (Fig. 2i, we have observed in populations of the lat j) and almost identical in terminal mor ter species. Records from the eastern Pacific phology to that herein figured for the ho region include mention of both Panopeus lotype of Acantholobulus schmitti (Fig. 2g, bermudensis (see Rathbun 1930, Garth h). However, as discussed below, the de scription of larval development for this spe 1961, Lemaitre 1981, Abele & Kim 1989, cies {Panopeus margentus) by Rodriguez & Lemaitre & Alvarez Leon 1992) and P. Spivak (2001) included some differences mirafloresensis (see Abele & Kim 1989: from that presented by Bakker et al. (1989) 36), and no attempt is herein made by us to for Hexapanopeus schmitti, which should resolve morphologically whether these east not be the case if these two names indeed ern Pacific populations could represent refer to the same species (further addressed more than one species. At least until further in Discussion below). comparative study is undertaken and on going molecular analyses are completed, Materials of Panopeus pacificus from we recognize A. mirafloresensis as the east Hawaii and Tahiti, along with some ques ern Pacific cognate of A. bermudensis. The tionably assigned to "Neopanope sp. ?" by natural range of latter species appears lim Edmondson (1931), were reviewed by For ited to warm-temperate and tropical shallow est & Guinot (1961) who provided detailed waters of the western Atlantic, as also con illustrations of gonopods, chelae, and cara cluded by Salgado Barragan & Hendriekx pace dentition for these specimens, all of (1997). which appear to be typical of Acantholob Systematic review of congeners.—Rec• ulus. Our direct examination of topotypic ognition of Panopeus margentus as a junior specimens of P. pacificus (four males, two synonym of Acantholobulus schmitti (Rath- females) collected by C. H. Edmondson bun, 1930), new combination, was based from Pearl Harbor (USNM 96304) confirms upon examination of the holotype (USNM this generic reassignment. We have also re 239191) and paratypes (USNM 239195) of examined the two males of P. pacificus re Panopeus margentus, all from Mar del Pla ported from Tahiti (MNHN B13963) by ta, Argentina. The description of P. mar Forest & Guinot (1961) and found no char gentus by Williams & Boschi (1990) in acters to distinguish them from the Hawai cluded no comparisons to or mention of ian materials. Edmondson (1931) preceded Hexapanopeus schmitti, even though the his original description of P. pacificus by latter species has been documented to range noting the possibility of this species being VOI.UME 116, NUMBER 2 447
"transported to Hawaii through shipments Panopeus mirafloresensis, 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 bermudensis, 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 mirafloresensis ent larger size evident in most specimens of A cantholobulus mirafloresensis, A. pacificus, there are also no obvious mor new combination phological characters to distinguish it from Panopeus margentus A. mirafloreaensis. 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 hemphillii genetic studies that we have in progress. junior synonym of A. bermudensis While we have not re-examined materials Hexapanopeus heblingi of the "Neopanope sp.?" 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 pacificus that we examined (USNM The use of brachyuran larval morpholo 96304) 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.?". In addition, the figure genetic questions is now relatively well es of the gonopod for "Neopanope sp.?" pro tablished (e.g., see references in Clark et al. vided by Forest & Guinot (1961: fig. 106) 1998). 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 superfamily Xanthoi- pected in gonopod morphology for species dea MacLeay, 1838 (e.g., Clark & Al-Ai- in this genus. We thus conclude that the pe daroos 1996, Clark Si Galil 1998, Clark & culiar cheliped dentition reported for "Neo Ng 1998, Fransozo et al. 2001, 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 are not uncommon Panopeidae Ortmann, 1893 sensu Guinot in populations of A. bermudensis that we 1978 (see also Martin & Davis 2001), lar have sampled in Florida (ULLZ collec val morphology is diverse. Prior to 1985, tions). These have also been illustrated for however, all known species of the Panopei variant specimens of both A. bermudensis dae save Panopeus bermudensis were (as Panopeus gatunensis, Abele & Kim, thought to have lateral carapace spines, 1989: fig. 15) and A. mirafloresensis, (as which is true of nearly all known xanthoid 448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON larvae. "Typical" zoeal larvae of panopeid Panopeus margentus with Acantholobulus crabs have well developed (although some schmitti, and conclusion that Hexapanopeus times short) lateral carapace spines, a ta hemphillii and H. heblingi must be synon- pering antennal protopod that is often ymized with Acantholobulus bermudensis. armed distally (and is never distally inflat In the case of Panopeus margentus, our rea ed), and two or three small spines on the sons for placing this species into synonymy telsonal furcae, all located at approximately with Acantholobulus schmitti on the basis the level of insertion of the three paired tel of adult features have been given earlier. sonal setae (Fig. 4). The larvae of P. ber One would thus expect that reported larval mudensis were described as differing from descriptions would prove to be identical, all other panopeid larvae in lacking these and to large extent this is true; Rodriguez lateral carapace spines (Martin et al. 1985), & Spivak (2001:818) state that "Panopeus having an antenna that is dilated at the tip margentus larvae are very similar to those and that lacks an exopod (actually present, of H. schmitti." The larvae of P. margentus but greatly reduced, see Discussion), and in as described by Rodriguez & Spivak (2001) bearing a single pair of telsonal spines lo also lack lateral carapace spines, although a cated posterior to the distal tip of the inte slight protrusion of the carapace can be rior telsonal setae. Remarkably, the zoeal seen (Fig. Ih, i), which may be a homo- stages of Hexapanopeus schmitti as de logue of the lateral spine in typical pano scribed by Bakker et al. (1989) also lack a peid zoeas. As described, however, the lateral carapace spine, have an antennal zoeal stages of P. margentus would appear protopod that is dilated at the tip, have a to lack the second pair of telson furcal reduced antennal exopod, and bear a pair of spines shown by Bakker et al. (1989) for posterior spines on the telsonal furca (al H, schmitti. though, in addition, there is also a pair of In the course of our studies, we have re more anterior furcal spines). examined the parental female (USNM Our decision to group the former Pano- 288046), as well as the larval stages of P. peus bermudensis and Hexapanopeus margentus (USNM 291175) described by schmitti, along with the former Panopeus Rodriguez & Spivak (2001). We cannot de mirafloresensis and P. pacificus, into Acan- finitively separate the female from those of tholobulus, new genus, on the basis of adult A. schmitti, though females of both of these morphology is thus supported by larval species remain poorly described. Among morphology for at least the first two of the larvae, when mounted in glycerin and these species. We expect that larval char examined with differential interference con acters of the former P. mirafloresensis and trast (DIC) optics, we do find zoeas at stag P. pacificus, which remain undescribed, es one through four in which a second, very should resemble those of the two other spe small pair of telson furcal spines can be dis cies of Acantholobulus. Given the similar cerned at high magnifications (first called to ities in adult morphology herewith reported our attention by J. Cuesta and B. Mahon). for proposed members of Acantholobulus, These, however, appear to be less conspic new genus, shared characters in larval mor uous than those illustrated for zoeal stages phology to the extent known within this of A. schmitti by Bakker et al. (1989), and group, and the established genetic distinc it should be noted that they were not ob tion for at least one of its members (Schu- vious on all specimens examined; in some bart et al. 2000), the generic revision herein cases, the small setal shaft appeared instead proposed is strongly supported. as a cluster of two or three miniscule setae Even so, some problems remain in our or setal fragments and may have been worn assignment of species to the new genus or damaged. Acantholobulus, decision to synonymize For Hexapanopeus heblingi we examined VOLUME 116, 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 bermudensis described by tions among the Xanthoidea. Martin et al. (1984, 1985). The absence of a lateral carapace spine, the inflated tip of Acknowledgments the antenna, the reduced antennal exopod, and other features are a nearly perfect We gratefully acknowledge the advice match, supporting our synonymy of the two and encouragement of the late R. B. Man species under the name Acantholobulus ning, who assisted us in early phases of this bermudensis. Yet, descriptions of the me- project. We also thank K. Anger, J. Cuesta, galopa appear to differ slightly. In the me- D. Guinot, R. Lemaitre, B. Mahon, F. Man- galopa of P. bermudensis, described by telatto, K. Reed, and C. Schubart, who in Martin et al. (1984, 1985), the frontal re various ways facilitated our completion of gion of the carapace bears anterolateral this paper. Support was provided from Na horns, and the cheliped bears a distinct re tional Science Foundation grants no. DEB curved spine on the ischium. Neither of 9972100 and DEB 9978193, U.S. Depart these characters is illustrated by Rodrigues ment of Energy grant no. DE-FG02- (1997) for the species therein referred to as 97ER12220, and several small grants from H. heblingi. Although frontal horns on the the Smithsonian Marine Station at Ft. carapace are easily missed, especially if the Pierce, Florida. This is contribution No. 94 carapace front is strongly deflected down of the UL Lafayette Laboratory for Crus ward, the cheliped spine is a feature that tacean Research and contribution No. 547 rarely escapes notice. of the Smithsonian Marine Station at Ft. 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., & W. Kim. 1986. An illustrated guide to and it is pertinent to note that Martin et al. the marine decapod crustaceans of Florida.— (1985) also overlooked a diagnostic struc State of Florida Department of Environmental ture in the course of describing larvae that Regulation Technical Series, Vol. 8, no. 1 (in 2 parts). 760 p. we now refer to Acantholobulus bermuden Abele, L. G., & W. Kim. 1989. 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 482:l-,'50. (ULLZ, uncatalogued slide mounts), and Bal
Department of Environmental Protection, 1998: first zoeal stage of Piatypodiella spectabilis i-xv, 1^238. (Herbst, 1794) (Decapoda, Brachyura, Xanthi Clark, R P., & A- M. Al-Aidaroos. 1996. The first zo- dae) obtained in the laboratory,—Gulf and Ca eas of Actaeodes hirsutissimus {Riippell, 1830) ribbean Research 13:79-85. and A. tomentosus (H. Milne Fxlwards. 1834) Fransozo, A., F, L. M. Mantelatto, & M, L. Negreiros- (Crustacea: Decapoda: Brachyura: Xanthidae: Fransozo. 1990. Larval development of Hexa- Actaeinae).—Journal of King Abdulaziz Uni panopeus paulensis Rathbun, 1930 (Crustacea, versity Marine Sciences 7, Special Issue: Sym Brachyura, Xanthidae) under laboratory condi posium on the Red Sea Marine Environment: tions,—Revista Brasileira Zoologia 7:31-45. 207-214. Garth, J. S. 1961. Part 2: Brachygnatha Brachyrhyn- Clark, R P., D. K. Calazans, & G. W. Pohle. 1998. cha. In Eastern Pacific expeditions of the New Accuracy and standardization of brachyuran lar York Zoological Society, XLV: Non-intertidal val descriptions.—Invertebrate Reproduction brachygnathous crabs from the west Coast of and Development 33:127 144. Tropical America.—Zoologica (New York) 46: Clark, R E, & B. S. Galil. 1998. The first stage zoea 133-159. 1 plate. of P.seudoliomera speciosa (Dana, 1852) (Crus Garth, J. S. 1991. Taxonomy, distribution, and ecology tacea, Decapoda, Brachyura, Xanthidae).— of Galapagos Brachyura. Pp. 123-145 in M, J. Zoosystema 20:193-200. James (ed.), Galapagos Marine Invertebrates,— Clark, R F, & R K. L, Ng. 1998. The larval devel Topics in Geobiology, vol. 8, Plenum Press, opment of the poisonous mosaic crab, Lopho- New York, z.ozymous pictor (Fabricius, 1798) (Crustacea, Gore, R. H., L. E. Scotto, & L, J, Becker. 1978. Com Decapoda, Brachyura, Xanthidae, Zosiminae), munity composition, stability, and trophic par with comments on familial characters for first titioning in decapod crustaceans inhabiting stage zoeas.—Zoosystema 20:201-220. some subtropical sabellariid worm reefs.—Bul Edmondson, C. H. 1931. New crustaceans from Kauai, letin of Marine Science 28:221-248. Oahu, and Maui. Bernice P. Bishop Museum Gore, R, H.. C. L. Van Dover. & K. A. Wilson. 1981. Occasional Papers 9(17): 1-18. Studies on decapod Crustacea from the Indian Felder, D. L. 1973. An annotated key to crabs and River region of Florida. XX. Micropanope bar- lobsters (Decapoda, Reptantia) from coastal wa badensis (Rathbun, 1921): the complete larval ters of the northwestern Gulf of Mexico.—Cen development under laboratory conditions ter for Wetland Resources, Louisiana State Uni (Brachyura, Xanthidae).—Journal of Crusta versity, Baton Rouge, 103 p. cean Biology 1:28-50. Felder, D. L., & A, H, Chaney. 1979. Decapod crus Guinot, D. 1967. Recherches preliminaires sur les tacean fauna of Seven and One-half Fathom groupements naturels chez les Crustaces Deca Reef, Texas: species composition, abundance, podes Brachyoures IIL A propos des affinites and species diversity,—Contributions in Marine des genres Dairoides Stebbing et Daira de Science 22:1-29, Haan.—Bulletin du Museum national d'Histoire Felder, D. L., J. W. Martin, & J. W. Goy. 1985. Patterns naturelle, 2e Serie 39:540-563. in early postlarval development of decapods. In Guinot, D, 1978. Principes d'une classification evolu A. M. Wenner (ed.). Larval Growth, vol. 2 of tive des Crustaces Decapodes Brachyoures.— F. Schram (ed.). Crustacean Issues. Pp. 163- Bulletin Biologique de le France et de la Bel- 225. Balkema Press, Rotterdam. gique (n. s.) 112:211-292. Ferreira, A. C, & C. Sankarankutty. 1997. Extension Lebour, M. 1944. Larval crabs from Bermuda.—Zool of the range of distribution of Panopeus mira- ogica (New York) 29:113-128. floresensis Abele & Kim (Crustacea: Decapoda: Lemaitre, R. 1981. Shallow-water crabs (Decapoda, Xanthidae).—Nauplius 5:155-157. Brachyura) collected in the southern Caribbean Forest, J., & D. Guinot. 1961. Crustaces Decapodes near Cartagena, Colombia,—Bulletin of Marine Brachyoures de Tahiti et des Tuamotu. Expe Science 31:234-266. dition Fran^aise sur les recifs coralliens de la Lemaitre, R.. & R. Alvarez Leon. 1992. Crustaceos Nouvelle-Cal^donie. Volume Preliminaire— decapodos del Pacifico colombiano: Lista de es- Editions de la Fondation Singer-Polignac, Paris: pecies y consideraciones zoogeograficas.—An- 1-195, 18 plates. ales del Instituto de Investigaciones Marinas, Fransozo, A. 1987. Desenvolvimento larval de Eriphia Punta Betin 21:.33-76. gonagra (Fabricius, 1781) (Decapoda, Xanthi Markham, J. C, & J. J. McDermott. 1981. A tabulation dae), em laboratorio.—Revista Brasileira Zool- of the Crustacea Decapoda of Bermuda. Pro ogia 4:165-179. ceedings of the Biological Society of Washing Fransozo, A., M. L. Negreiros-Fransozo, J. W. Martin, ton 83(4); 1266-1276. & S. E. Trautwein. 2001. Morphology of the Martin, J. W. 1984. Notes and bibliography on the lar- VOLUME 116, NUMBER 2 451
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