Review of the Western Atlantic Porcellanidae (Crustacea: Decapoda:
Anomura) with New Records, Systematic Observations, and
Comments on Biogeography
IRENE TERESA RODR?GUEZ1, GONZALO HERN?NDEZ2, AND DARRYL L. FELDER1*
1Department of Biology, University of Louisiana at Lafayette, P.O. Box 42451, Lafayette, Louisiana 70504, U.S.A.
*Corresponding author: dlf4517@louisiana.edu
2Universidad de Oriente, Apartado Postal 074, La Asunci?n, Isla Margarita, Venezuela
ABSTRACT.?Fifty years following a major revision by Haig (1956) and after numerous additions and
changes of status, the porcelain crab fauna of the western Atlantic numbers 48 valid species plus two
undescribed species herein reported (Neopisosoma sp. and Pachycheles sp.). Eleven genera are known for the
region, including three new genera (Neopisosoma, Clastotoechus, and Parapetrolisthes). The monospecific
genus Madarateuchus, established by Harvey (1999) to receive Clastotoechus vanderhorsti, is rejected. Pisidia
was resurrected, and Porcellanopsis was combined with Megalobrachium. Prompted by our efforts to under-
stand biogeographic relationships of porcellanids from Bocas del Toro, Panama, our study of over 400
western Atlantic records revealed undescribed species and northward range extensions for four species
(Petrolisthes quadratus to U.S.A., Petrolisthes rosariensis and P. sanmartini to Belize, and Megalobrachium
mortenseni to Honduras). Preliminary morphological and molecular studies have shown that Petrolisthes
armatus, P. galathinus, and P. tridentatus present considerable intra-specific variability throughout their
range, suggesting that further revisionary studies are required to fully resolve their taxonomic status.
KEYWORDS.?Porcellanids, porcelain crabs, Gulf of Mexico, distribution
INTRODUCTION
Porcellanidae Haworth, 1825, is a family
of crab-shaped anomuran crustaceans com-
monly known as porcelain crabs. This fam-
ily is considered a well-defined taxon
within the superfamily Galatheoidea
Samouelle, 1819 (see Haig 1960). Other
members of the superfamily are Aeglidae
Dana, 1852, Chirostylidae Ortmann, 1892,
and Galatheidae Samouelle, 1819 (Martin
and Davis 2001). Porcelain crabs number
about 280 species partitioned among 30
genera. These crabs occur worldwide in
primarily intertidal and sublittoral zones of
tropical and subtropical regions, and oc-
cupy habitats such as depressions under
stones, spaces in worm tubes, cavities of
sponges, and excavations in coral reefs.
Most species are free-living but a few live
commensally (Haig 1960; Gore 1972b, 1982;
Harvey 1999; Hern?ndez 1999; Werding
1983b; Werding et al. 2003).
Nearly half of the species in this family
have been recorded in the Americas (Gore
1982). Seven species are reported to occur
on both the Atlantic and the Pacific coasts:
Pachycheles chacei Haig, 1956, P. monilifer
(Dana, 1852), Petrolisthes armatus (Gibbes,
1850), P. galathinus (Bosc, 1802), P. robsonae
Glassell, 1945, P. tonsorius Haig, 1960, and
P. tridentatus Stimpson, 1859 (see Haig
1956, 1960; Gore and Abele 1974; Scelzo
1982; Abele and Kim 1989; Hiller et al.
2004).
In the almost fifty years since the major
revision of the Porcellanidae from the west-
ern North Atlantic by Haig (1956), impor-
tant efforts have been made to better un-
derstand the composition, diversity,
distribution, and relationships of these
crabs in the western Atlantic (see Boschi
1963; Co?lho 1964, 1966; Rodrigues da
Costa 1964, 1968; Rickner 1975b; Haig 1962,
1966; Gore 1970a, 1974, 1982, 1983; Gore
and Abele 1974, 1976; Werding 1977, 1978a,
1978b, 1982, 1983a, 1984, 1986, 1996; Rod-
r?guez 1980; Scelzo 1982, 1983; Silva et al.
Caribbean Journal of Science, Vol. 41, No. 3, 544-582, 2005
Copyright 2005 College of Arts and Sciences
University of Puerto Rico, Mayagu?ez
544
T
A
B
L
E
1.
St
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PORCELLANIDAE (CRUSTACEA) 545
1989; Veloso and Melo 1993; Hern?ndez
and Bola?os 1995; Hern?ndez 1999; Le-
maitre and Campos 2000; Werding and
Hiller 2002; Werding et al. 2001; Werding
and Kraus 2002; Werding et al. 2003). The
morphology of larval and megalopal stages
of porcelain crabs is very useful in elucidat-
ing relationships among species. Ontoge-
netic studies are currently available for 27
species distributed in the western Atlantic
(Table 1).
In addition to systematics, some progress
has been made in ecological (e.g., Gray
1961; Gore and Shoup 1968; Sandifer 1973;
Caine 1975; Markham 1975; Felder and
Chaney 1979; Wenner and Read 1982;
Young 1986; Schoppe and Werding 1996;
Stillman and Somero 2000) and physiologi-
cal fields (Teytaud 1971; Craig 1974; L?pez-
Greco et al. 2002). However, no compre-
hensive synthesis of the taxonomic
literature of this group of anomurans is
available for the western Atlantic. This re-
view provides that synthesis and updates
records since publication of Haig?s (1956)
monograph. It was undertaken as an essen-
tial step in ongoing molecular (Rodr?guez
et al. in review) and morphological studies
of porcellanid crabs from the vicinity of Bo-
cas del Toro, Panama, and adjacent western
Atlantic waters.
Porcelain crabs numbered 29 species for
the western North Atlantic at the time of
Haig?s (1956) compilation. Pachycheles gree-
leyi (Rathbun, 1900) and P. laevidactylus Ort-
mann, 1892, from Brazil were not included
in that report. Taking these two species into
account, the known porcellanid fauna of
the western Atlantic was 31 species at that
time. In the following decades, a number of
additional species were described, and one
species, Petrolisthes robsonae, previously
known only from the eastern Pacific, was
reported from the Caribbean (Haig 1960;
Abele and Kim 1989). Some species have
been recognized as synonyms, others as in-
valid, and many range extensions have
been documented for the region. An up-
dated list of 48 valid species and their dis-
tributional patterns was recently published
(Werding et al. 2003).
As evident from literature, studies of
porcellanids along mainland coastlines of
Central America bordering the Caribbean
Sea, especially in areas adjacent to Panama,
are scarce; similarly, few works address
specifically this fauna further north and
within the Gulf of Mexico (Haig 1956; Rick-
ner 1975b; Werding et al. 2003). Thorough
compilation of Gulf of Mexico records and
systematic re-examinations of materials
upon which those records are based has
never been undertaken, leaving much
question as to their relationships with pu-
tative Caribbean conspecifics. In this study
we focused on extensive new collections
not only from the Caribbean coast of
Panama, but also materials from the Gulf of
Mexico and adjacent waters of Florida,
U.S.A., Quintana Roo, Mexico, Belize, Hon-
duras, and Venezuela. In addition, we in-
cluded Pacific records of amphi-American
species.
MATERIALS AND METHODS
The synonymy for each genus is in-
cluded in this review, with the generic type
species shown in parenthesis. The status of
each genus is discussed, followed by infor-
mation for each species. Synonymies for
species include references to the original
description and type locality, followed by
references subsequent to Haig?s (1956) re-
port.
Localities are listed in geographical or-
der from north to south, west to east, and
by country. In the western Atlantic, these
are listed by zoogeographical provinces as
follows: Virginian, from Cape Cod to
Cape Hatteras; northern Warm-temperate
(= Carolinian), from Cape Hatteras to Cape
Canaveral; Gulf of Mexico Warm-temper-
ate, from Cape Romano to Cape Rojo,
Mexico; Caribbean, from Cape Rojo to the
Orinoco Delta; West Indian, includes the
Caribbean Islands; Brazilian, from the
Orinoco Delta to Cape Frio; and Argentin-
ian, from Cape Frio to 43/44?S (modified
after Briggs 1974; Williams 1984; Boschi
2000). Pacific localities are treated at the
end of accounts for materials examined.
Additional information to support each rec-
ord is provided in Appendix I (available at:
www.caribjsci.org/dataarchive.html.
I. T. RODR?GUEZ ET AL.546
The majority of the material examined is
part of the Zoological Collection of the Uni-
versity of Louisiana at Lafayette (ULLZ),
where records resulted from 37 years of
collecting effort, dating back to 1967. In the
last 5 years, the number of these holdings
doubled as the result of major field expedi-
tions targeting coastal porcelain crabs and
other decapods, including particularly pro-
ductive efforts in the vicinity of Bocas del
Toro on the Caribbean coast of Panama.
Other primary sites for these collecting
campaigns included Fort Pierce, Florida
(July 2000, July-August 2001), southern
Texas (September 2001), U.S.A.; Veracruz
to Quintana Roo, Mexico (July 2002); Belize
(October 2002); Pacific coast of Nicaragua
(September 2000, November 2001); and
Baja California, Mexico (December 2001).
Specimens from Venezuela are also in-
cluded in the study (March-June 1996,
April-June, 2001, October 2004). The Colec-
ci?n Nacional de Crust?ceos, Instituto de
Biolog?a, Universidad Nacional Aut?noma
de M?xico, M?xico, D. F., M?xico (CNCR)
and the National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D. C., U.S.A. (USNM), provided additional
material for study. In addition to the rec-
ords for the western Atlantic, those for the
eastern Pacific are included for amphi-
American species.
In the distributional accounts, localities
are treated in geographic order as noted
above. Those in the western Atlantic are
cited first followed in succession by those
in the eastern Pacific and those from the
eastern Atlantic. No attempt was made to
comprehensively include every mention of
a species? name to be found in theses, dis-
sertations, and meeting abstracts. We also
did not exhaustively review listings in in-
structional manuals, keys lacking original
illustrations or records, and textbooks. In
those few cases where records have been
based upon theses or abstracts, the speci-
men identifications were independently
verified by one or more of the present au-
thors.
Color photographs of selected Panama-
nian specimens featured herein were ob-
tained in the field by posing freshly chilled
specimens, submerged beneath water, un-
der illuminantion from direct and reflected
sunlight. Digital photographs were ob-
tained with a 35 mm Fuji Finepix camera
equipped with a Nikon 60 mm Micro
Nikkor macrolens. Sex and carapace width
(? 0.01 mm = cw) of each photographed
specimen were determined prior to archi-
val in the University of Louisiana Zoologi-
cal Collections (ULLZ).
RESULTS
The porcellanid collection at ULLZ rep-
resents 467 records that include over 2,300
specimens belonging to more than 70 spe-
cies. Sixteen countries and three oceanic re-
gions are represented: the Indo-Pacific with
5 records, eastern Pacific with 75 records,
and western Atlantic with 387 records. The
western Atlantic component contains 206
records for the Gulf of Mexico. The follow-
ing systematic account incorporates all the
records for the western Atlantic, but only
the amphi-American records of the eastern
Pacific. Thirteen records from the CNCR
and one from the USNM are also included.
Systematic Account
Family Porcellanidae Haworth, 1825
Genus Clastotoechus Haig, 1960
Clastotoechus Haig, 1960:175-176 (type
species Petrolisthes diffractus Haig, 1957, by
original designation).?Harvey, 1999:4-7.?
Rodr?guez et al., in review.
Madarateuchus Harvey, 1999:27 (type spe-
cies Clastotoechus vanderhorsti, by mono-
typy).?Rodr?guez et al., in review.
Remarks.?The genus Clastotoechus was
proposed to include two western Atlantic
species, Petrolisthes nodosus Streets, 1872,
and P. vanderhorsti Schmitt, 1924, and one
eastern Pacific species, P. diffractus Haig,
1957. Three more species of the genus were
subsequently described: C. gorgonensis
Werding and Haig, 1983, C. hickmani Har-
vey, 1999, and C. lasios Harvey, 1999, all
from the eastern Pacific.
The monotypic genus Madarateuchus
Harvey, 1999, was originally proposed to
receive Clastotoechus vanderhorsti and dis-
tinguish this species from the rest of the
PORCELLANIDAE (CRUSTACEA) 547
species in the genus Clastotoechus. How-
ever, this species shares unique characters
with species of Clastotoechus that clearly set
them apart from other genera within the
family (see Haig 1960:175). Moreover, in re-
cent studies based on molecular data, C.
nodosus and C. vanderhorsti were clearly
grouped closely together in a clade sepa-
rated from other genera of the family
(Werding et al. 2001, on the basis of partial
DNA-sequences of COI gene; Rodr?guez
et al. in review, on the basis of partial
sequences of the 16S rRNA gene). Further
molecular studies including more species
of the genus, should better clarify their
relationships with C. vanderhorsti. Given
this information and pending ongoing
studies by Werding et al. (Werding,
personal communication), recognition of
the genus Madarateuchus is not justified
and it is herein synonymized with Clastoto-
echus.
Clastotoechus nodosus (Streets, 1872)
Petrolisthes nodosus Streets, 1872:133 (type
locality St. Martin Island).?Haig, 1956:27-
28.?Rodr?guez, 1980:218.
Clastotoechus nodosus.?Haig, 1960:177,
fig. 5(3).?Rickner, 1975b:163.?Gore and
Abele, 1976:15, fig. 2.?Werding, 1977:178,
fig. 3 (not fig. 2).?Scelzo, 1982:1131.?
Alvarez et al., 1999:9.?Harvey, 1999:7, figs.
2, 4-15.?Hern?ndez, 1999:249, table 2.?
Werding et al., 2001:108-109, table 3, fig.
2.?Werding et al., 2003:79-85, tables 1, 2.?
Hern?ndez et al., 2003:419-428, figs. 1-6.?
Galicia-Castillo and Herna?ndez-Aguilera,
2005:239, 241, fig. 2.?Rodr?guez et al., in
review.
Clastoechus nodosus.?Werding, 1978b:
215, figs. 1a, b (misspelling).
Clastoechus modosus.?Werding, 1978b:
220 (misspelling).
Material examined.?Punta Delgada
(ULLZ 4092, 5233, 5367 to 5369), El Morro
(ULLZ 5231, 5232), La Mancha (ULLZ 5370,
5371), Escondida Beach (ULLZ 5372), Vera-
cruz, Mexico. Charagato Bay, Cubagua Is-
land, Venezuela (ULLZ 5341).
Distribution.?Veracruz, Mexico; Galeta
Island, Panama; Santa Marta, Colombia;
Cura?ao; La Tortuga and Cubagua Is-
lands, Venezuela; Cuba; Puerto Rico; St.
Martin.
Clastotoechus vanderhorsti (Schmitt, 1924)
Petrolisthes vanderhorsti Schmitt, 1924:73,
pl. 8, fig. 7 (type locality Caracas Bay,
Cura?ao).?Haig, 1956:27.?Rodr?guez,
1980:218.
Clastotoechus vanderhorsti.?Haig, 1960:
175, fig. 5 (2).?Lewis, 1960:426.?Haig,
1962:184.?Teytaud, 1971:43.?Markham,
1975:266.?Werding, 1977:180, fig. 2 (not
fig. 3).?Scelzo, 1982:1131.?Werding,
1983b:3.?Schoppe, 1991:373.?Schoppe,
1994:107-119, figs. 1-8.?Schoppe and
Werding, 1996:181-186.?Hern?ndez, 1999:
246, 249, tables 1, 2.?Werding et al., 2001:
108-109, table 3, fig. 2.?Werding et al.,
2003:79-85, tables 1, 2.?Rodr?guez et al., in
review.
Clastoechus vanderhorsti.?Werding,
1978b:215, 220 (misspelling).
Madarateuchus vanderhorsti.?Harvey,
1999:27, figs. 64-75.
Material examined.?La Tortuga Island,
Venezuela (ULLZ 5342).
Distribution.?Gulf of Urab?, Santa
Marta, and Guajira Colombia; Cura?ao and
Bonaire; La Tortuga and Cubagua Islands,
Venezuela; Cuba; St. Croix, Virgin Islands;
Saba; Martinique; Barbados.
Genus Euceramus Stimpson, 1860
Euceramus Stimpson, 1860:445 (type spe-
cies Euceramus praelongus Stimpson, 1860,
by monotypy).?Haig, 1960:187-188.?
Rodr?guez et al., in review.
Remarks.?Euceramus occurs on both
coasts of the Americas. E. praelongus Stimp-
son, 1860, is the only representative of the
genus in the western Atlantic, where it is
restricted to the northern temperate region
in the Virginian and Warm-temperate
Provinces (modified after Williams 1984).
The species is remarkably rare (Haig 1956),
being a highly adapted burrower in sub-
tidal sandy and rubble-covered beaches
(Williams 1984). In the eastern Pacific the
genus is represented by two species, E. pan-
atelus Glassell, 1938, and E. transversilinea-
tus (Lockington, 1878) (see Haig 1960).
I. T. RODR?GUEZ ET AL.548
Euceramus praelongus Stimpson, 1860
Euceramus praelongus Stimpson, 1860:
444-445 (type locality Beaufort, North Caro-
lina).?Haig, 1956:7.?Haig, 1960:187-
190.?Tabb and Manning, 1961:599.?
Williams, 1965:109, fig. 86.?Roberts, 1968:
121-130, figs. 1-3, table 1.?Rouse, 1970:
140.?Gosner, 1971:539-540, fig. 21-54B.?
Menzel, 1971:78.?Van Engel and Sandifer,
1972:157.?Felder, 1973:32, pl. 4, fig. 11.?
Sandifer, 1973:245.?Camp et al., 1977:29.?
Young, 1978:176.?Maris, 1983:237-246,
figs. 1-7.?Truesdale and Andryszak, 1983:
43, table 2.?Williams, 1984:239, fig. 174.?
Abele and Kim, 1986:410, 424, 425, fig. a.?
Ruppert and Fox, 1988:251, 404.?Britton
and Morton, 1989:121, fig. 6-7 D, 245-246,
fig 10-1 L, 258.?Williams et al., 1989:35.?
Tunnell and Alvarado, 1996:136.?Camp et
al., 1998:144.?Hern?ndez, 1999:246, 249,
tables 1, 2.?Werding et al., 2003:79-85,
table 1.?Galicia-Castillo and Herna?ndez
Aguilera, 2005:237.
Material examined.?Fort Pierce, Florida
(ULLZ 5245, 5806), offshore Louisiana
(ULLZ 4575), Heald Banks, offshore Texas
(ULLZ 4457), northern Gulf of Mexico
(ULLZ 1308, 1309), U.S.A.
Distribution.?Chesapeake Bay North
Carolina; Charleston, South Carolina; east-
ern and western Florida; Louisiana; and
Texas; U.S.A.
Remarks.?The species is endemic to the
Virginian and northern Warm-temperate
Provinces in the western Atlantic.
Genus Megalobrachium Stimpson, 1858
Megalobrachium Stimpson, 1858:228 (type
species Megalobrachium granuliferum Stimp-
son, 1858, by original designation).?Haig,
1960:212-213.?Rodr?guez et al., in review.
Porcellanides Nobili, 1901:21 (type species
Porcellanides festae Nobili, 1901, by mono-
typy).
Porcellanopsis Rathbun, 1910:601 (type
species Porcellanides festae Nobili, 1901, by
monotypy; Porcellanides preoccupied by
Porce l lanides Czerniavsky, 1884) .?
Rodr?guez et al., in review.
Pisonella Glassell, 1938:436.
Remarks.?The genus Megalobrachium
contains 12 species distributed on both
coasts of the Americas. Haig (1956) re-
ported M. poeyi (Gu?rin, 1855) as the only
representative of this genus in the western
North Atlantic, but included two species of
Porcellanopsis Rathbun, 1910, P. rosea (Rath-
bun, 1900) and P. soriata (Say, 1818). Porcel-
lanopsis was subsequently combined with
Megalobrachium (Haig, 1960). Two more
species were later described, M. mortenseni
Haig, 1962, and M. walteri Rodrigues da
Costa, 1968. The latter has been shown to
be a synonym of M. soriatum (Veloso and
Melo 1993). Four species of this genus are
now recognized for the western Atlantic.
Preliminary molecular analyses (Rodr?guez
et al. in review) suggest the validity of Por-
cellanopsis and the need for further study.
Megalobrachium mortenseni Haig, 1962
Plate 1, fig. A
Megalobrachium mortenseni Haig, 1962:
189, figs. 2-5 (type locality west of Congo
Cay, northwest of St. John, Virgin Is-
lands).?Gore, 1970a:965.?Co?lho, 1971:
233.?Co?lho and Ramos, 1972:175.?
Werding, 1977:181, fig. 4.?Werding, 1978b:
215.?Corredor et al., 1979:32.?Gore, 1982:
6.?Scelzo, 1982:1131.?Werding, 1982:
441.?Veloso and Melo, 1993:175.?
Hern?ndez, 1999:249, table 2.?Melo, 1999:
218-219, figs. 141, 142.?Lira et al., 2001:55,
56, 62-64, fig. 4.?Werding et al., 2003:79-85,
tables 1, 2.
Material examined.?Caribe Point, Roat?n,
Bay Islands, Honduras (ULLZ 5450). Bocas
del Toro, Panama (ULLZ 5998, 6005, 6007,
6088).
Distribution.?Roat?n, Bay Islands, Hon-
duras; Bocas del Toro and Lim?n Bay,
Panama; Santa Marta, Colombia; Margarita
Island, Venezuela; St. John, Virgin Islands;
from Par? to Sao Paulo, Brazil.
Remarks.?The species is first recorded in
Honduras and its range is extended north-
ward. Two morphotypes have been ob-
served among specimens from Panama. In
one, the dorsal surface of the carpus and
manus has longitudinal rows of pits and is
covered with small, coarse granules, as
originally described by Haig (1962). In the
other, those longitudinal rows of pits can-
not be clearly distinguished, and the dorsal
surface of the carpus and manus is uneven
PORCELLANIDAE (CRUSTACEA) 549
and more heavily granulated (see Plate 1,
fig. A).
Megalobrachium poeyi (Gu?rin, 1855)
Plate 1, fig. B
Porcellana poeyi Gu?rin, 1855, pl. 2, fig. 4
(type locality Cuba).
Megalobrachium poeyi.?Haig, 1956:33-34
[part, Atlantic only].?Haig, 1960:214-215
[part, Atlantic only].?Haig, 1962:188-189
[part, Atlantic only].?Haig, 1968:57, 71-72
[part, Atlantic only].?Co?lho, 1966:63
[part, Atlantic only].?Co?lho, 1971:233.?
Gore, 1971a:404-425, figs. 2, 4, 6, 8, 10, 11C,
PLATE 1. Selected porcellanids from the vicinity of Bocas del Toro, Panama. A. Megalobrachium mortenseni
Haig, 1962, female, cw = 2.60 mm, ULLZ 6088. B. Megalobrachium poeyi (Gu?rin, 1855), female, cw = 4.90 mm,
ULLZ 6094. C. Megalobrachium roseum (Rathbun, 1900), female, cw = 3.46 mm, ULLZ 6095. D. Pachycheles chacei
Haig, 1956, male, cw = 2.46 mm, ULLZ 6096. E. Pachycheles cristobalensis Gore, 1970, male, cw = 4.24 mm, ULLZ
6097. F. Pachycheles serratus (Benedict, 1901), male, cw = 5.40 mm, ULLZ 6098. G. Petrolisthes caribensis Werding,
1983, male, cw = 4.00 mm, ULLZ 5997. H. Petrolisthes galathinus (Bosc, 1802), female, cw = 11.32 mm, ULLZ 6087.
I. Petrolisthes jugosus Streets, 1872, female, cw = 3.90 mm, ULLZ 6106.
I. T. RODR?GUEZ ET AL.550
Fi Pl t 1 li 4/C
F, tables 1-3 [part, Atlantic only].?Gore,
1972a:67-90, figs. 13-18, table 5 [part, Atlan-
tic only].?Gore, 1974:704.?Gore and
Abele, 1974:559, 568-573, fig. E.?Gore
and Abele, 1976:17.?Voss, 1976:92-
93, fig.?Camp et al., 1977:29.?Werding,
1977:182, figs. 1b, 5.?Werding, 1978b:
215.?Rodr?guez, 1980:219.?Gore, 1982:
6.?Scelzo, 1982:1131.?Werding, 1984:5.?
Abele and Kim, 1986:412, 414, 415, fig. b.?
Williams et al., 1989:35.?Camp et al., 1998:
144.?Hern?ndez, 1999:246, 249, tables 1,
2.?Lira et al., 2001:55, 56, 58-60, fig. 2.?
Werding et al., 2003:79-85, tables 1, 2.?
Galicia-Castillo and Herna?ndez-Aguilera,
2005:239, 243-244, fig. 5.?Rodr?guez et al.,
in review.
Material examined.?Indian River, Fort
Pierce, Florida, U.S.A. (ULLZ 3610).
Chemuyil, Quintana Roo, Mexico (CNCR
9818). Bocas del Toro, Panama (ULLZ
6094). Bar?, Colombia (ULLZ 4885). El
Manglillo, Macanao Peninsula, Margarita
Island, Venezuela (ULLZ 5343).
Distribution.?Eastern Florida, U.S.A.;
Quintana Roo, Mexico; Bocas del Toro and
Caledonia Bay, Panama; Old Providence Is-
land and continental coast of Colombia;
Coche, Margarita, and Cubagua Islands,
Venezuela; Greater and Lesser Antilles;
Paraiba and Sao Paulo, Brazil.
Remarks.?Prior to restriction of this spe-
cies to Atlantic representatives, references
to it commonly encompassed its Pacific sib-
ling, which is now recognized as Megalobra-
chium pacificum Gore and Abele, 1974.
Megalobrachium roseum (Rathbun, 1900)
Plate 1, fig. C
Porcellana rosea Rathbun, 1900:148, pl. 8,
fig. 3 (type locality Mamanguape, Brazil).
Porcellanopsis rosea.?Haig, 1956:34-35.?
Co?lho, 1966:59.?Rodr?guez, 1980:219.
Megalobrachium roseum.?Haig, 1960:225,
227.?Haig, 1966:356.?Co?lho, 1971:233.?
Co?lho and Ramos, 1972:175.?Scelzo and
Boschi, 1973:213.?Gore and Abele, 1976:
17.?Werding, 1977:183, fig. 6.?Werding,
1978b:217.?Gore, 1982:7.?Scelzo, 1982:
1131.?Young, 1986:103.?Veloso and
Melo, 1993:175.?Hern?ndez, 1999:249,
table 2.?Melo, 1999:220-221, figs. 143,
144.?Lira et al., 2001:55, 56, 60-62, fig. 3.?
Hern?ndez et al., 2002:113-125, figs. 1-6.?
Werding et al., 2003:79-85, tables 1, 2.
Material examined.?Bocas del Toro,
Panama (ULLZ 6095, 6101). El Manglillo,
Macanao Peninsula, Margarita Island, Ven-
ezuela (ULLZ 5344).
Distribution.?Bocas del Toro, Panama;
Santa Marta, Colombia; Margarita and
Cubagua Islands, La Restringa Lagoon,
Venezuela; Martinique; from Maranhao to
Sao Paulo, Brazil.
Megalobrachium soriatum (Say, 1818)
Porcellana soriata Say, 1818:456 (type lo-
cality St. Catherine?s Island, Georgia).?
Pounds, 1961:28.?Leary, 1964:28.
Porcellanopsis soriata.?Haig, 1956:35-
36.?Tabb and Manning, 1961:599.?
Co?lho, 1964:255.?Bullis and Thomp-
son, 1965:10.?Co?lho, 1966:60.?Menzel,
1971:79.
Megalobrachium soriatum.?Haig, 1960:
229.?Williams, 1965:112, fig. 89.?Haig,
1966:356.?McCloskey, 1970:30.?Rouse,
1970:141.?Co?lho and Ramos, 1972:175.?
Felder, 1973:32-35, pl. 4, fig. 12.?Gore,
1973b:837-856, figs. 1-5, tables 1, 2.?Gore
and Abele, 1976:17, fig. 3.?Werding, 1977:
185, fig. 7.?Gore et al., 1978:225.?Young,
1978:176.?Felder and Chaney, 1979:25, 28,
fig. 3 table 1, appendices I, II.?Hern?ndez-
Aguilera and Sosa-Hern?ndez, 1982:42-
43.?Reed et al., 1982:768.?Scelzo, 1982:
1131.?Wenner and Read, 1982:187, table
2.?Maris, 1983:237-246, figs. 1-7.?
Williams, 1984:240, fig. 175.?Abele and
Kim, 1986:412, 414, 415, fig. a.?Young,
1986:103.?Ruppert and Fox, 1988:249,
404.?Williams et al., 1989:35.?Markham
et al., 1990:427.?Veloso and Melo, 1993:
175.?Hern?ndez, 1999:246, 249, tables 1,
2.?Melo, 1999, p. 222-223, figs. 145, 146.?
Hern?ndez-Aguilera et al., 1996:49.?
Tunnell and Alvarado, 1996:136.?Camp et
al., 1998:144.?Alvarez et al., 1999:9.?Lira
et al., 2001:55, 56-68, fig. 1.?Werding et al.,
2003:79-85, tables 1, 2.?Galicia-Castillo
and Herna?ndez-Aguilera, 2005:239, 244-
245, fig. 6.?Rodr?guez et al., in review.
Megalobrachium walteri Rodrigues da
Costa, 1968.?Veloso and Melo, 1993:175.
Material examined.?Fort Pierce (ULLZ
5964) and Bahia Honda State Park (ULLZ
PORCELLANIDAE (CRUSTACEA) 551
5262), Florida; San Padre Island (ULLZ
2167, 5279) and 71?2 Fathom Reef (ULLZ
2151 to 2166, 2168, 2169), Texas, U.S.A.
Barra del Tordo, Tamaulipas (ULLZ 5819);
Reef off Puerto Morelos, Quintana Roo
(CNCR 5734), Mexico. Bocas del Toro,
Panama (ULLZ 6002, 6086).
Distribution.?North Carolina, South
Carolina, Georgia, eastern and western
Florida, Texas, U.S.A.; Tamaulipas, Vera-
cruz, Campeche, and Quintana, Mexico;
Bocas del Toro and Bah?a Caledonia,
Panama; Santa Marta, Colombia; Margarita
Island, Venezuela; Martinique; Barbados;
from Cear? to Sao Paulo, Brazil.
Remarks.?While preliminary genetic
analyses suggest that trans-Floridian popu-
lations of Megalobrachium soriatum are ge-
netically divergent (Rodr?guez et al. in re-
view), differences in morphology were not
readily apparent between specimens from
eastern Florida (Bahia Honda, Florida
Keys) to those from the Gulf of Mexico.
Genus Minyocerus Stimpson, 1858
Minyocerus Stimpson, 1858:229 (type spe-
cies Porcellana angusta Dana, 1852, by origi-
nal designation).?Haig, 1960:193.?
Rodr?guez et al., in review.
Porcellina M?ller, 1862:194 (type species
Porcellina stellicola M?ller, 1862, by mono-
typy).
Remarks.?Minyocerus is represented by
only two species, M. angustus (Dana, 1852)
in the western Atlantic and M. kirki Glas-
sell, 1938 in the eastern Pacific. Both species
seem to be obligatory commensals on echi-
noderms, mainly confined to species of the
genus Luidia (Asteroidea) and also found
on a few genera of Ophiuroidea (Haig 1956,
1960; Werding 1983b).
Minyocerus angustus (Dana, 1852)
Porcellana angusta Dana, 1852:423 (type
locality Rio de Janeiro, Brazil).
Minyocerus angustus.?Haig, 1956:30-
31.?Holthuis, 1959:161.?Haig, 1962:
184.?Rodrigues da Costa, 1964:565.?
Haig, 1966:354.?Co?lho, 1966:61.?Fausto-
Filho, 1967:12.?Gore and Shoup, 1968:240-
248, figs. 1-3.?Gore, 1970a:967.?Co?lho,
1971:233.?Co?lho and Ramos, 1972:174.?
Gore, 1974:705.?Werding, 1977:187, fig.
8.?Rodr?guez, 1980:218.?Scelzo, 1982:
1132.?Silva et al., 1989:138, figs. 8, 14.?
Veloso and Melo, 1993:176.?Hern?ndez,
Bola?os, and Graterol, 1996:87-92, figs.
1-3.?Hern?ndez, 1999:246, 249, tables 1,
2.?Melo, 1999:224-225, figs. 147, 148.?
Werding et al., 2003:79-85, tables 1, 2.?
Hern?ndez et al., in press.
Material examined.?Punta Arenas, Cuba-
gua Island, Venezuela (ULLZ 5966).
Distribution.?Belize; Honduras; Nicara-
gua; Costa Rica; Lim?n Bay and Col?n,
Panama; Gulf of Morosquillo and Santa
Marta, Colombia; La Tortuga, Margarita,
and Cubagua Islands, Venezuela; Suri-
name; from Par? to Santa Catarina, Brazil.
Genus Neopisosoma Haig, 1960
Neopisosoma Haig, 1960:123-124 (type
species Neopisosoma bicapillatum Haig, 1960,
by original designation).?Werding et al.,
2001:105-110, tables 1-3.?Rodr?guez et al.,
2004:299, 306-307.?Rodr?guez et al., in re-
view.
Remarks.?Neopisosoma was proposed to
contain a group of species from the two
coasts of the Americas, which were previ-
ously grouped in the genera Pisosoma
Stimpson, 1858, and Pachycheles Stimpson,
1858. In addition to the two species recog-
nized by Haig (1956), N. angustifrons (Bene-
dict, 1901) and N. curacaoense Schmitt, 1924,
Werding (1986) described two other spe-
cies, N. neglectum Werding, 1986 and N. ori-
entale Werding, 1986, from the Caribbean.
An undescribed species from Quintana
Roo, Mexico, was found in the course of the
present study and is being described at this
time. The genus contains four valid species
in the western Atlantic and three in the
eastern Pacific, in addition to at least one
more western Atlantic species currently in
description.
Two morphologically different groups of
Neopisosoma can be distinguished (see Haig
1960; Werding et al. 2001; Rodr?guez et al.
2004). One group is characterized by the
subquadrate carapace; side walls incom-
plete, with a narrow projection extending
to mesobranchial level; chelipeds with flat-
tened granules and carpus with crests; tel-
I. T. RODR?GUEZ ET AL.552
son 5 or 7-plated; and male pleopods pres-
ent. This group includes N. bicapillatum, N.
curacaoense, N. mexicanum (Streets, 1871), N.
neglectum, and N. orientale. The second
group has a rounded carapace; side walls
incomplete, ending at the epibranchial
level; chelipeds with rounded, projecting
granules and no crests on carpus; telson
5-plated; no male pleopods. This group
contains N. angustifrons, N. dohenyi Haig,
1960, and Neopisosoma sp. Additionally, the
paraphyly of Neopisosoma was suggested by
a molecular analysis (Werding et al. 2001).
The status of Neopisosoma as a distinct ge-
nus from Pachycheles was discussed by
Gore (1977), Werding (1986), Werding and
M?ller (1990), Werding et al. (2001), and
Rodr?guez et al. (2004).
Neopisosoma angustifrons (Benedict, 1901)
Pisosoma angustifrons Benedict, 1901:135,
pl. 3, fig. 6 (type locality Trinidad).?Haig,
1956:15.
Neopisosoma angustifrons.?Haig, 1960:
124, 131.?Haig, 1962:181.?Rickner, 1975b:
163.?Gore and Abele, 1976:18.?Gore,
1977:258-300, figs. 1-8.?Werding, 1977:187,
fig. 9.?Werding, 1978b:217.?Rodr?guez,
1980:214, pl. 2.?Scelzo, 1982:1132.?Abele
and Kim, 1986:410, 424, 425, fig. b.?
Werding, 1986:162, fig. 1.?Williams et al.,
1989:35.?Hern?ndez-Aguilera et al., 1996:
49.?Camp et al., 1998:144.?Alvarez et al.,
1999:9.?Hern?ndez, 1999:246, 249, tables 1,
2.?Werding et al., 2001:105-110, fig. 2,
tables 1-3.?Werding et al., 2003:79-85,
tables 1, 2.?Rodr?guez et al., 2004:301, 306,
table 3.?Galicia-Castillo and Herna?ndez-
Aguilera, 2005:240, 242-243.?Rodr?guez et
al., in review.
Material examined.?Punta Delgada
(ULLZ 4091, 5235, 5373 to 5384, 5451), El
Morro (ULLZ 5234), La Mancha (ULLZ
5385, 5386), Escondida Beach (ULLZ 5387),
Veracruz, Mexico. Charagato Bay, Cubagua
Island, Venezuela (ULLZ 5345).
Distribution.?Veracruz, Mexico; Galeta
Island, Panama; Santa Marta, Colombia;
Cura?ao and Bonaire; La Tortuga, Mar-
garita, and Cubagua Islands, Venezuela;
Trinidad; Bahamas; Puerto Rico; Saba;
Guadeloupe.
Neopisosoma curacaoense (Schmitt, 1924)
Pisosoma curacaoensis Schmitt, 1924:75, pl.
8, figs. 1-3 (type locality Spanish Bay,
Cura?ao).
Pisosoma curacaoense.?Haig, 1956:15-16.
Neopisosoma curacaoense.?Haig, 1960:
124, 126-127.?Haig, 1962:181.?Rodr?guez,
1980:214.?Scelzo, 1982:1132.?Werding,
1986:162, 166, figs. 2a-j.?Markham et al.,
1990:427.?Hern?ndez, 1999:249, table 2.?
Werding et al., 2001:107, table 1.?Werding
et al., 2003:79-85, tables 1, 2.?Galicia-
Castillo and Herna?ndez-Aguilera, 2005:240,
242-243.
Neopisosoma caracaoensis.?Galicia-
Castillo and Herna?ndez-Aguilera, 2005:
243, fig. 4 (misspelling).
Not Neopisosoma curacaoense.?Werding,
1978b:213-214, 217-218, 220, figs. 2a, b.
Not N. curacaoensis.?Werding, 1978b:
214.
Material examined.?None.
Distribution.?Quintana Roo, Mexico;
Aruba, Cura?ao, and Bonaire; La Tortuga
Island, Venezuela; Bahamas; Puerto Rico;
St. Martin; Virgin Islands; Saba; Grenada.
Remarks.?The record for Quintana Roo,
Mexico (Markham et al. 1990) was based on
one specimen and no further material has
been reported from the area. Markham et
al. (1990) included Colombia in the distri-
bution of N. curacaoense, based on the
record presented in Werding (1978); how-
ever, this record was later described as N.
neglectum Werding, 1986, thus excluding N.
curacaoense from Colombia (see Werding
1986). Galicia-Castillo and Herna?ndez-
Aguilera (2005:243) in error listed Colom-
bia as part of the range of N. curacaoense.
Neopisosoma neglectum Werding, 1986
Neopisosoma curacaoense.?Werding,
1978b:213-214, 217-218, 220, figs. 2a, b.
Neopisosoma neglectum Werding, 1986:
162, 169, fig. 3 (type locality Santa Marta,
Colombia).?Werding and M?ller, 1990:
363-374, figs. 1-6.?Hern?ndez, 1999:246,
249, tables 1, 2.?Werding et al., 2001:105-
110, tables 1-3, figs. 1, 2.?Werding et al.,
2003:79-85, tables 1, 2.
Material examined.?None.
PORCELLANIDAE (CRUSTACEA) 553
Distribution.?Gulf of Dari?n, Gulf of
Urab?, and Santa Marta, Colombia; St. Mar-
tin; Grenada.
Neopisosoma orientale Werding, 1986
Neopisosoma orientale Werding, 1986:162,
172, fig. 4 (type locality Blanchisseuse Bay,
Trinidad).?Lira, 1997:57, fig. 15.?
Hern?ndez, 1999:249, table 2.?Werding et
al., 2001:107, fig. 2, table 1.?Werding et al.,
2003:79-85, tables 1, 2.
Material examined.?La Pared, Macanao
Peninsula, Margarita Island, Venezuela
(ULLZ 5346).
Distribution.?Margarita Island, Venezu-
ela; Blanchisseuse Bay, Trinidad.
Neopisosoma sp.
Material examined.?Punta Nizuc, Can-
c?n, Quintana Roo, Mexico (ULLZ 4844).
Remarks.?This is a new species currently
being described by D. L. Felder and I. T.
Rodr?guez.
Genus Pachycheles Stimpson, 1858
Pachycheles Stimpson, 1858:228 (type spe-
cies Porcellana grossimana Gu?rin, 1835, by
original designation).?Haig, 1960:131-
132.?Rodr?guez et al., in review.
Pisosoma Stimpson, 1858:228 (type spe-
cies Porcellana pisum H. Milne Edwards,
1837, by original designation).?Rodr?guez
et al., in review.
Remarks.?The genus Pachycheles is the
second most speciose of the family and in-
cludes over 40 valid species worldwide dis-
tributed. Haig (1956) included seven spe-
cies in her list, leaving out the Brazilian
endemic P. greeleyi (Rathbun, 1900) and P.
laevidactylus Ortmann, 1892. These species
were initially described from Brazil and
later recognized as valid species by Harvey
and De Santo (1996), who also considered
P. haigae Rodrigues da Costa, 1960, a junior
synonym of P. laevidactylus. Subsequent to
Haig?s paper, three additional species have
been described: P. chubutensis Boschi, 1963,
a southern temperate species, P. cristobalen-
sis Gore, 1970a, and P. susanae Gore and
Abele, 1974, the latter two from the Car-
ibbean Sea. A new species from Bocas del
Toro, Panama, was found in this study and
is being described. To date, there are 12
western Atlantic valid species in this genus.
On the basis of molecular characters of the
mitochondrial genes COI (Werding et al.
2001) and 16S rRNA (Stillman and Reeb
2001; Rodr?guez et al. in review), the genus
appears to be paraphyletic and is in need of
revision.
Pachycheles ackleianus A. Milne-Edwards,
1880
Pachycheles ackleianus A. Milne-Edwards,
1880:36 (type localities west coast of
Florida, U.S.A. and Jolbos Islands,
Mexico).?Haig, 1956:13.?Haig, 1960:143-
144.?Haig, 1962:182.?Co?lho, 1964:255.?
Bullis and Thompson, 1965:10.?Co?lho,
1966:52.?Haig, 1966:352.?Gore, 1970a:
962.?Co?lho, 1971:233.?Co?lho and Ra-
mos, 1972:173.?Gore, 1974:705, fig. 2.?
Rickner, 1975b:162.?Werding, 1977:188,
fig. 10.?Corredor et al., 1979:32.?
Westinga and Hoetjes, 1981:141.?Gore,
1982:7.?Scelzo, 1982:1132.?Werding,
1982:441.?Werding, 1983b:5.?Abele and
Kim, 1986:412, 416, 417, fig. d.?Williams et
al., 1989:35.?Markham et al., 1990:427.?
Veloso and Melo, 1993:176.?Camp et al.,
1998:144.?Hern?ndez, 1999:249, table 2.?
Melo, 1999:228-229, figs. 149, 150.?
Werding et al., 2003:79-85, tables 1, 2.?
Rodr?guez et al., 2004:291.?Galicia-Castillo
and Herna?ndez-Aguilera, 2005:239, 245-
246, fig. 7.?Rodr?guez et al., in review.
Material examined.?Looe Key Reef,
Florida (ULLZ 4824, 4826), off Tortugas,
Florida (ULLZ 5798), offshore western
Florida (ULLZ 5278), off Sanibel Island,
western Florida, (ULLZ 5669), U.S.A. Roca
Chica, Macanao Peninsula, Margarita Is-
land, Venezuela (ULLZ 5347).
Distribution.?Tampa Bay and Florida
Keys, Florida, U.S.A.; Veracruz, Yucat?n,
and Quintana Roo, Mexico; Mulatas Archi-
pelago and Canal Zone, Panama; Cabo de
La Vela and Santa Marta, Colombia; Los
Roques, Margarita, and Los Testigos Is-
lands, Venezuela; Cuba; Jamaica; Domini-
can Republic; St. Thomas; Antigua; Marti-
nique; St. Lucia; Barbados; Guyanas;
Pernambuco to Bahia, Brazil.
I. T. RODR?GUEZ ET AL.554
Pachycheles chacei Haig, 1956
Plate 1, fig. D
Pachycheles chacei Haig, 1956:9, pl. 1, figs.
1 -6 ( type loca l i ty Caledonia Bay ,
Panama).?Haig, 1960:134, pl. 31, fig. 3.?
Haig, 1968:57, 68.?Gore and Abele, 1976:
19.?Werding, 1977:190, fig. 11.?Werding,
1978b:217.?Corredor et al., 1979:32.?
Gore, 1982:8.?Scelzo, 1982:1133.?
Werding, 1982:441.?Werding, 1984:5.?
Lemaitre and Alvarez-Le?n, 1992:47, table
1.?Hern?ndez and Bola?os, 1995:75-76.?
Hern?ndez, 1999:249, table 2.?Hern?ndez
et al., 1999:27-29, fig. 2, table 1.?Melo,
1999:230-231, figs. 151, 152.?Werding et
al., 2001:107, table 1.?Werding et al., 2003:
79-85, tables 1, 2.?Rodr?guez et al., 2004:
291.
Material examined.?Bocas del Toro,
Panama (ULLZ 6096, 6102). Rosario Is-
lands, Colombia (ULLZ 5294).
Distribution.?Western Atlantic: Bocas
del Toro and Gulf of Dari?n, Panama, to
Santa Marta, Colombia; Old Providence
and Rosario Islands, Colombia; Margarita
Island, Venezuela; Alag?as, Brazil. Eastern
Pacific: San Jos?, Guatemala; Acajutla, El
Salvador; Verde Island and Pi?as Bay,
Panama; Humboldt Bay, Colombia; La Li-
bertad and Santa Elena Bay, Ecuador.
Pachycheles chubutensis Boschi, 1963
Pachycheles chubutensis Boschi, 1963:35,
figs. 2, 3a, e, g (type locality Gulf of San
Mat?as, Chubut, Argentina).?Haig, 1966:
353.?Co?lho and Ramos, 1972:173.?
Scelzo and Boschi, 1973:207, table 1.?
Boschi, 1979:137.?Bremec and Cazzaniga,
1984:149-162, figs. 3, 4.?Silva et al., 1989:
135, figs. 3, 4, 17-21.?Veloso and Melo,
1993:177.?Hern?ndez, 1999:249, table 2.?
Melo, 1999:232-233, figs. 153, 154.?
Werding et al., 2003:79-85, table 1.
Material examined.?None.
Distribution.?From Florian?polis, Brazil,
to Chubut, Argentina.
Pachycheles cristobalensis Gore, 1970
Plate 1, fig. E
Pachycheles cristobalensis Gore, 1970a:958,
figs. 1, 2 (type locality Lim?n Bay,
Panama).?Gore and Abele, 1976:19.?
Werding, 1977:175.?Gore, 1982:8.?Scelzo,
1982:1133.?Hern?ndez, 1999:249, table
2.?Werding et al., 2003:79-85, tables 1, 2.?
Rodr?guez et al., 2004:291.
Material examined.?Bocas del Toro,
Panama (ULLZ 6097, 6103).
Distribution.?Lim?n Bay, Bocas del
Toro, and Galeta Island Panama.
Remarks.?This species is endemic to the
Atlantic coast of Panama.
Pachycheles greeleyi (Rathbun, 1900)
Pisosoma greeleyi Rathbun, 1900:174, pl. 8,
fig. 4 (type locality Coral Reef, Maceio, Ala-
g?as, Brazil).
Pachycheles greeleyi.?Haig, 1956:9.?
Haig, 1966:352.?Co?lho, 1966:52.?Co?lho,
1971:233.?Co?lho and Ramos, 1972:173.?
Young, 1986:103.?Veloso and Melo, 1993:
177.?Hern?ndez, 1999:249, table 2.?Melo,
1999:234-235, figs. 155, 156.?Werding et
al., 2003:79-85, tables 1, 2.
Pachycheles greelei.?Werding et al., 2001:
107, table 1 (misspelling).
Material examined.?None.
Distribution.?From Par? to Esp?rito
Santo, Brazil.
Remarks.?The species is endemic to Bra-
zil.
Pachycheles laevidactylus Ortmann, 1892
Pachycheles laevidactylus Ortmann, 1892:
266, pl. 12, fig. 1 (type locality Brazil; this
locality is incorrect fide Ortmann, 1897:293
in Haig, 1960:167).?Harvey and DeSanto,
1996:710, fig. 2.?Melo, 1999:236-237, figs.
157, 158.?Werding et al., 2003:79-85, tables
1, 2.?Rodr?gue?z et al., 2004:291, 302-305,
tables 1, 2.
Pachycheles grossimanus Ortmann, 1897:
292.?Haig, 1955:43 (in part).?Haig, 1960:
167, pl. 35, fig. 1 (in part).
Pachycheles haigae Rodrigues da Costa,
1960:21, figs. 1-4.?Boschi, 1963:31, figs. 1,
3.?Haig, 1966:353.?Boschi et al., 1967:6-
26, figs. 1-8, table 1.?Co?lho and Ramos,
1972:173.?Scelzo and Boschi, 1973:207,
table 1.?Boschi, 1979:137.?Bremec and
Cazzaniga, 1984:149-162, fig. 2.?Silva et
al., 1989:134, figs. 2, 11.?Boschi et al., 1992:
PORCELLANIDAE (CRUSTACEA) 555
56.?Veloso and Melo, 1993:177.?
Hern?ndez, 1999:246, 249, tables 1, 2.
Pachycheles rudis Moreira, 1901:32, 91.
Pachycheles haige.?Co?lho, 1966:54 (mis-
spelling).
Material examined.?Rio de Janeiro, Brazil
(ULLZ 5439).
Distribution.?From Pernambuco to
Santa Catarina, Brazil; Rocha, Uruguay;
Mar de Plata and Miramar, Argentina.
Pachycheles monilifer (Dana, 1852)
Porcellana rugosimanus White, 1847:63
(nomen nudum).
Porcellana monilifera Dana, 1852:413; 1855,
pl. 26, fig. 3 (type locality Rio de Janeiro,
Brazil).
Pachycheles monilifer.?Rathbun, 1900:
148.?Haig, 1956:13.?Rodriguez, 1959:
274.?Haig, 1960:160-162, pl. 33, fig. 4.?
Haig, 1962:183.?Rodrigues da Costa, 1962:
5, 7.?Rodrigues da Costa, 1964:565.?
Co?lho, 1966:53.?Haig, 1966:353.?Co?lho
and Ramos, 1972:172.?Gore, 1973a:132-
150, figs. 1-6, table 1.?Gore, 1974:707.?
Rickner, 1975b:162.?Camp et al., 1977:
29.?Werding, 1977:190, fig. 12.?Gore et
al., 1978:225.?Rodr?guez, 1980:213, pl. 1.?
Reed et al., 1982:768.?Scelzo, 1982:1133.?
Abele and Kim, 1986:412, 418, 419, fig. a.?
Silva et al., 1989:133, figs. 1, 10.?Williams
et al., 1989:35.?Veloso and Melo, 1993:
178.?Hern?ndez-Aguilera et al., 1996:49.?
Camp et al., 1998:144.?Alvarez et al., 1999:
9.?Hern?ndez, 1999:246, 249, tables 1, 2.?
Hern?ndez et al., 1999:27, table 1.?Melo,
1999:238-239, figs. 159, 160.?Werding et
al., 2003:79-85, tables 1, 2.?Rodr?guez et
al., 2004:291, tables 1, 2.?Hiller et al., 2004:
130, 134, 135, table 1.?Galicia-Castillo and
Herna?ndez-Aguilera, 2005:239, 246-247, fig.
8.?Rodr?guez et al., in review.
Material examined.?Fort Pierce (ULLZ
5246), Jenson Beach (ULLZ 3631), off St. Lu-
cie (ULLZ 5823), eastern Florida; Atlantic
coast of Florida (ULLZ 5644, 5657, 5956,
5957), U.S.A. Barra del Tordo, Tamaulipas
(ULLZ 5821); Punta Delgada (ULLZ 5236),
Escondida Beach (ULLZ 5388), Veracruz,
Mexico. Charagato Bay, Cubagua Island,
Venezuela (ULLZ 5348).
Distribution.?Western Atlantic: Eastern
Florida, U.S.A.; Tamaulipas, Veracruz,
Campeche, and Quintana Roo, Mexico;
Santa Marta, Cabo de la Vela, and Rosario
Islands, Colombia; Margarita and Cubagua
Islands, Venezuela; Trinidad; St. Thomas
and St. John; from Par? to Santa Catarina,
Brazil. Eastern Pacific: La Libertad, Ecua-
dor; Cancas, Peru.
Remarks.?Haig (1960) stated that her rec-
ord of Pachycheles monilifer for the eastern
Pacific was based on a single juvenile fe-
male and that it was tentatively referred to
this species, but the exact determination of
its status must await recovery of adult ma-
terial from the Pacific coast. New records
recently confirmed the presence of this spe-
cies in the eastern Pacific (Hiller et al. 2004).
Pachycheles pilosus (H. Milne Edwards,
1837)
Porcellana pilosa H. Milne Edwards, 1837:
225 (type locality Charleston, South Caro-
lina).
Pachycheles pilosus.?Schmitt, 1924:76.?
Haig, 1956:11.?Haig, 1962:182.?Williams,
1965:108, fig. 84.?Werding, 1977:192, fig.
13.?Young, 1978:176.?Corredor et al.,
1979:32.?Rodr?guez, 1980:214.?Scelzo,
1982:1133.?Werding, 1982:441.?Werding,
1984:6.?Markham et al., 1990:427.?
Williams 1984:241, fig. 176.?Abele and
Kim, 1986:412, 416, 417, fig. a.?Scelzo and
Varela, 1988:39, fig. 2.?Williams et al.,
1989:35.?Mart?nez-Iglesias et al., 1993:
13.?Hern?ndez-Aguilera et al., 1996:49.?
Camp et al., 1998:144.?Garc?a et al., 1998:
28-29, table 1.?Hern?ndez, 1999:249, table
2.?Nizenski, 2003:116.?Werding et al.,
2003:79-85, tables 1, 2.?Rodr?guez et al.,
2004:291.?Galicia-Castillo and Herna?n-
dez-Aguilera, 2005:239, 247, fig. 9.?Pi?ate
et al., 2005.?Rodr?guez et al., in review.
Material examined.?Canc?n (ULLZ
4845), Puerto Morelos (CNCR 5645), Maja-
hual (ULLZ 5389), Quintana Roo, Mexico.
Cayo de Indios, off Isle of Pines (= Isla de la
Juventud), Cuba (ULLZ 5454). Punta Brava,
Rosario Islands, Colombia (ULLZ 4483). La
Tortuga Island, Venezuela (ULLZ 5349).
Distribution.?Charleston, South Caro-
lina, Key Biscayne and Key West, eastern
I. T. RODR?GUEZ ET AL.556
Florida, Sarasota Bay, western Florida,
U.S.A.; Campeche and Quintana Roo,
Mexico; Cartagena, Santa Marta, Old Provi-
dence and Rosario Islands, Colombia;
Aruba, Cura?ao, and Bonaire; Aves, La
Blanquilla, and La Tortuga Islands, Ven-
ezuela; Tobago; Bahamas; Cuba; Puerto
Rico; St. Thomas; Guadeloupe; Barbados.
Pachycheles riisei (Stimpson, 1859)
Pisosoma riisei Stimpson, 1858:228 (nomen
nudum), 1859:75 (type locality St. Thomas).
Pachycheles riisei.?Haig, 1956:12.?Haig,
1962:182.?Co?lho, 1966:54.?Co?lho and
Ramos, 1972:172.?Fausto-Filho, 1974:8.?
Werding, 1977:192, fig. 14.?Werding,
1978b:219.?Corredor et al., 1979:32.?
Scelzo, 1982:1133.?Werding, 1982:441.?
Werding, 1984:6.?Abele and Kim, 1986:
412, 416, 417, fig. b.?Young, 1986:103.?
Williams et al., 1989:35.?Veloso and Melo,
1993:178.?Hern?ndez and Bola?os, 1995:
7 5 - 7 6 . ? C a m p e t a l . , 1 9 9 8 : 1 4 4 . ?
Hern?ndez, 1999:249, table 2.?Melo, 1999:
240-241, figs. 161, 162.?Werding et al.,
2001:105-106, fig. 1.?Werding et al., 2003:
79-85, tables 1, 2.?Rodr?guez et al., 2004:
291.
Material examined.?Carrie Bow Cay, Be-
lize (ULLZ 5453). El Manglecito, Cubagua
Island, Venezuela (ULLZ 5350).
Distribution.?Key West, Florida, U.S.A.;
Carrie Bow Cay, Belize; Gulf of Dari?n,
Santa Marta, Old Providence and Rosario
Islands, Colombia; Margarita and Cubagua
Islands, Venezuela; Tobago; Puerto Rico;
St. Thomas; Barbados; Fernando de No-
ronha, Trindade, Para?ba and Alag?as, Bra-
zil.
Remarks.?The first record for Belize is
presented in this study.
Pachycheles rugimanus
A. Milne-Edwards, 1880
Pachycheles rugimanus A. Milne-Edwards,
1880:36 (type localities Contoy and west of
Florida).?Haig, 1956:12.?Co?lho, 1964:
255.?Williams, 1965:108, fig. 85.?Cain,
1972:80.?Young, 1978:176.?Soto, 1980:
90.?Scelzo, 1982:1133.?Wenner and Read,
1982:186, 195, tables 2, 5.?Williams, 1984:
242, fig. 177.?Abele and Kim, 1986:412,
416, 417, fig. c.?Williams et al., 1989:35.?
Markham et al., 1990:427.?Veloso and
Melo, 1993:178.?Camp et al., 1998:144.?
Alvarez et al., 1999:9.?Hern?ndez, 1999:
249, table 2.?Melo, 1999:242-243, figs. 163,
164.?Werding et al., 2001:105-106, fig. 1.?
Nizenski, 2003:117.?Werding et al., 2003:
79-85, tables 1, 2.?Galicia-Castillo and
Herna?ndez-Aguilera, 2005:239, 247-248, fig.
10.
Material examined.?Off Florida (ULLZ
2043, 2048), off Tampa Bay, Florida (ULLZ
5824), U.S.A.
Distribution.?Beaufort and Cape Look-
out, North Carolina and west Florida,
U.S.A.; Veracruz and Quintana Roo,
Mexico; St. Thomas, Virgin Islands; Suri-
name; from Amap? to Pernambuco, Brazil.
Pachycheles serratus (Benedict, 1901)
Plate 1, fig. F
Pisosoma serrata Benedict, 1901:135, pl. 3,
fig. 7 (type locality Mayagu?ez, Puerto Rico).
Pachycheles serratus.?Haig, 1956:8-9.?
Haig, 1962:181.?Gore, 1970a:962, fig. 2.?
Gore and Abele, 1976:20.?Werding, 1977:
194, fig. 15.?Werding, 1978b:219.?
Corredor et al., 1979:32.?Gore, 1982:9.?
Scelzo, 1982:1133.?Werding, 1982:441.?
Scelzo, 1983:80.?Hern?ndez, 1999:246, 249,
tables 1, 2.?Werding et al., 2001:107-109,
fig. 2, tables 1, 3.?Lira et al., 2003:149-
151.?Werding et al., 2003:79-85, tables 1,
2.?Rodr?guez et al., 2004:291-301, figs. 1-6,
tables 1, 2.
Material examined.?Bocas del Toro,
Panama (ULLZ 6001, 6098, 6104, 6108).
Punta Bol?var, Colombia (ULLZ 5295).
Distribution.?Bocas del Toro and Cale-
donia Bay, Panama; Santa Marta and Punta
Bol?var, Colombia; Margarita Island, Ven-
ezuela; Puerto Rico; St. Thomas.
Pachycheles susanae Gore and Abele, 1974
Pachycheles susanae Gore and Abele, 1974:
560, fig. 1 (type locality Galeta Island,
Panama).?Gore and Abele, 1976:20.?
Werding, 1977:195, fig. 16.?Werding,
1978b:219.?Corredor et al., 1979:32.?
Gore, 1982:10.?Scelzo, 1982:1133.?
Werding, 1982:442.?Scelzo, 1983:80.?
Scelzo, 1984:374.?Hern?ndez, 1999:249,
PORCELLANIDAE (CRUSTACEA) 557
table 2.?Werding et al., 2001:105-106, fig.
1, table 1.?Werding et al., 2003:79-85,
tables 1, 2.?Rodr?guez et al., 2004:291.?
Galicia-Castillo and Herna?ndez-Aguilera,
2005:237, 239, 248-249, fig. 11.?Rodr?guez
et al., in review.
Material examined.?Puerto Morelos,
Quintana Roo, Mexico (CNCR 7273). Bocas
del Toro (ULLZ 6089) and Mar?a Grande
(ULLZ 5351), Panama. El Manglecito,
Cubagua Island, Venezuela (ULLZ 5352).
Distribution.?Quintana Roo, Mexico; Bo-
cas del Toro, Mar?a Grande, and Galeta Is-
land, Panama; Rosario Islands and Santa
Marta, Colombia; Isla Cubagua, Venezuela;
Martinique.
Pachycheles sp.
Material examined.?Bocas del Toro,
Panama (ULLZ 6107).
Distribution.?Bocas del Toro, Panama.
Remarks.?This is a new species being de-
scribed by D. L. Felder and I. T. Rodr?guez.
Genus Parapetrolisthes Haig, 1962
Parapetrolisthes Haig, 1962:173 (type spe-
cies Petrolisthes tortugensis Glassell, 1945, by
monotypy).?Rodr?guez et al., in review.
Remarks.?The genus Parapetrolisthes
Haig, 1962, was defined for a species pre-
viously included in the genus Petrolisthes
Stimpson, 1858. P. tortugensis shows a num-
ber of characters not present in any other
porcellanid species. The peculiar spoon-
shaped finger tips and the compressed,
truncate teeth that occur on the dactylus of
the major cheliped have been considered
homologous to those in the family Gala-
theidae (H. Kraus et al. unpublished).
However, molecular analyses (partial se-
quences of the 16S rRNA gene) group this
species within the clade of the P. galathinus-
complex, supporting their close relation-
ship (Rodr?guez et al. in review). The pres-
ence of distinct, transverse, piliferous
striations on the carapace seems to be a uni-
fying character within the complex, while
the shape of the fingers and chelipeds in
Parapetrolisthes, appears to reflect feeding
adaptations convergent with some gala-
theid forms.
Parapetrolisthes tortugensis (Glassell, 1945)
Petrolisthes tortugensis Glassell, 1945:228,
fig. 2 (type locality Tortugas, Florida).?
Haig, 1956:22.?Rodr?guez, 1980:217.
Parapetrolisthes tortugensis.?Haig, 1962:
173, fig. 1.?Gore, 1974:707.?Corredor et
al., 1979:32.?Scelzo, 1982:1131.?Werding,
1982:442, fig. 2.?Lemaitre, 1984:428.?
Abele and Kim, 1986:410, 424, 425, fig. c.?
Williams et al., 1989:35.?Camp et al., 1998:
144.?Hern?ndez, 1999:249, table 2.?
Werding et al., 2003:79-85, tables 1, 2.?
Rodr?guez et al., in review.
Material examined.?Looe Key Reef,
Florida Keys, U.S.A. (ULLZ 4825, 5825).
Distribution.?Looe Key Reef and Dry
Tortugas, Florida, U.S.A.; Rosario Islands
and Santa Marta, Colombia; La Tortuga Is-
land, Venezuela; Bahamas; Virgin Islands.
Genus Petrolisthes Stimpson, 1858
Petrolisthes Stimpson, 1858:227 (type spe-
cies Porcellana violacea Gu?rin, 1831, by
original designation).?Haig, 1960:21-24.?
Rodr?guez et al., in review.
Remarks.?Petrolisthes is the most speci-
ose of all porcellanid genera with over 100
valid species worldwide. Haig (1956) rec-
ognized ten species for the western North
Atlantic and later classified two of them in
the genus Clastotoechus Haig, 1960, and one
in the genus Parapetrolisthes Haig, 1962. In
the latter paper, she reincorporated Petrolis-
thes jugosus Streets, 1872, into the genus
Petrolisthes, where it was initially included
before being treated by Haig (1956) as Piso-
soma jugosum. In the following decades
eight new species from the western Atlantic
were assigned to the genus (Gore 1983;
Werding 1978a, 1978b, 1983a, 1996; Wer-
ding and Hiller 2002; Werding and Kraus
2002). Petrolisthes tonsorius Haig, 1960, for-
merly known from only the eastern Pacific,
was reported from the southern Caribbean
Sea (Werding 1977; Hern?ndez et al. 1999),
and P. robsonae Glassell, 1945, another Pa-
cific species, apparently migrated through
the Panama Canal and was reported sev-
eral times on the Atlantic side of the Canal
(Haig 1960; Abele and Kim 1989). Petrolis-
thes cessacii A. Milne-Edwards, 1878, from
I. T. RODR?GUEZ ET AL.558
the Gulf of Mexico and the Caribbean, was
found to be a junior synonym of P. margi-
natus Stimpson, 1859 (see Gore 1982, 1983),
and P. serratus Henderson, 1888 from Bra-
zil, was concluded to be a junior synonym
of P. amoenus (Gu?rin, 1855) (Rathbun
1900). Petrolisthes costai Rodrigues da Costa,
1968, was never confirmed as a valid spe-
cies. To date, the genus includes 18 valid
species in the western Atlantic.
Petrolisthes amoenus (Gu?rin, 1855)
Porcellana amoena Gu?rin, 1855, pl. 2, fig.
2 (type locality Cuba).
Petrolisthes amoenus.?Haig, 1956:25-
26.?Chace, 1956a:152.?Haig, 1962:177.?
Rodrigues da Costa, 1964:565.?Haig, 1966:
352.?Co?lho, 1971:233.?Co?lho and
Ramos, 1972:173.?Gore, 1974:707, fig. 3.?
Werding, 1977:198, fig. 17.?Rodr?guez,
1980:217.?Scelzo, 1982:1133.?Werding,
1984:7, fig. 1.?Veloso and Melo, 1993:
179.?Hern?ndez, 1999:249, table 2.?Melo,
1999:246-247, figs. 165, 166.?Werding et
al., 2003:79-85, tables 1, 2.?Galicia-Castillo
and Herna?ndez-Aguilera, 2005:240, 249,
fig. 12.
Petrolisthes serratus Henderson, 1888:107,
pl. 11, figs. 2, 2a.?Co?lho, 1966:58.?
Fausto-Filho, 1975:80.
Petrolisthes? serratus.?Fausto-Filho,
1974:8.
Material examined.?El Manglecito, Cuba-
gua Island, Venezuela (ULLZ 5353).
Distribution.?Florida Straits, U.S.A.; Yu-
cata?n, Mexico; Old Providence Island and
Santa Marta, Colombia; Cura?ao and Bo-
naire; Los Roques and Cubagua Islands,
Venezuela; Trinidad; Cuba; Puerto Rico;
Barbados; Alag?as and Bahia, Brazil.
Petrolisthes armatus (Gibbes, 1850)
Porcellana armata Gibbes, 1850:190 (type
locality Florida).
Petrolisthes armatus.?Haig, 1956:19.?
Chace, 1956b:14, 17-21, fig. 5.?Haig, 1959:
328.?Parker, 1959:2132, 2135, 2161, pl. IV,
fig. 22a, b, appendix table II.?Haig, 1960:
25, 50-55, 57, 59, 269-272, 386-387, pl. 19, fig.
2, table 11.?Tabb and Manning, 1961:
599.?Haig, 1962:178.?Leary, 1964:27-
28.?Bullis and Thompson, 1965:10.?Haig,
1968:57, 62.?Co?lho, 1966:55.?Westervelt,
1967:34-37, 129, fig. 10.?Fausto-Filho, 1970:
58.?Gore, 1970a:964.?Gore, 1970b:75-89,
figs. 1-6.?Rouse, 1970:140.?Co?lho, 1971:
233.?Co?lho and Ramos, 1972:174.?Gore,
1972a:12-66, figs. 1-12, tables 1-3.?Gore,
1972b:67-83, figs. 1-6, tables 1, 2.?Brusca,
1973: 229-230, fig. 7.32.?Felder, 1973:33?
35, pl. 4, fig. 13.?Fausto-Filho, 1974:8.?
Gore, 1974:709.?Fotheringham and
Brunenmeister, 1975:17, 28, 138, 167, fig.
2.14.?Gore and Abele, 1976:21.?Camp et
al., 1977:30.?Werding, 1977:198, fig. 18.?
Werding, 1978b:219.?Corredor et al., 1979:
32.?Brusca, 1980:262, 264, 266, pl. 2, fig.
17.1.?Carvacho, 1980:250, table 1.?
Rodr?guez, 1980:215.?Markham and Mc-
Dermott, 1981:1271.?Gore, 1982:11.?
Scelzo, 1982:1134.?Werding, 1982:442.?
Coen and Heck, 1983:213.?Maris, 1983:
237-246, figs. 1-7.?Abele and Kim, 1986:
413, 420, 421, fig. c.?Chace et al., 1986:338,
pl. 10.3, 10.4.?Raz Guzm?n et al., 1986:
348.?Campos-Gonz?lez and Mac?as-
Ch?vez, 1987: 241-244, table 1.?Abele and
Kim, 1989:21.?Britton and Morton, 1989:
45-46, 68, 80, 82, 90-91, figs. 3-2 H, 4-3 C, 5-1
M.?Camp et al., 1998:144.?Silva et al.,
1989:136, figs. 5, 16.?Williams et al., 1989:
35.?Markham et al., 1990:427.?Mart?nez-
Iglesias and Alcolado, 1990:table 26.?
Human, 1992:164-165.?Lemaitre and
Alvarez Le?n, 1992:48, table 1.?Veloso and
Melo, 1993:179.?Hern?ndez-Alvarez,
1995:35-36, pl. 7, fig. 1.?Osawa, 1995:167,
t a b l e 5 . ? G r a t e r o l , 1 9 9 6 : 1 - 1 0 0 . ?
Hern?ndez-Aguilera et al., 1996:50.?
Tunnell and Alvarado, 1996:136.?Alvarez
et al., 1999:9.?Hern?ndez, 1999:246, 249,
tables 1, 2.?Hern?ndez et al., 1999:27, table
1.?Melo, 1999:248-249, figs. 167, 168.?
Knott et al., 2000:404.?Stillman and Som-
ero, 2000:200-208, figs. 1-4.?Villalobos,
2000:184, 186-188, pl. 4, fig. 1, table 5.?
Diaz-Ferguson and Vargas-Zamora, 2001:
97-101.?Stillman and Reeb, 2001:236-243,
tables 1-3, figs. 1, 2.?Nizenski, 2003:117.?
Werding et al., 2003:79-85, tables 1, 2.?
Galicia-Castillo and Herna?ndez-Aguilera,
2005:240, 249-250, fig. 13.?Garcia-
Guerrero et al., 2005:339, 347, 348, table 1.?
Rodr?guez et al., in review.
Material examined.?Western Atlantic:
PORCELLANIDAE (CRUSTACEA) 559
Wilmington, North Carolina (ULLZ 4247);
Sea Island, Georgia (ULLZ 5261); Fort
Pierce, Florida (ULLZ 3646, 4484, 4656,
4848, 5247 to 5253), Hutchinson Island,
Florida (ULLZ 3767), Atlantic coast of
Florida (ULLZ 5656, 5903), Ambersands
Beach, Sebastian, Florida (ULLZ 5959),
Looe Key, Florida (ULLZ 5263), Everglades
City, Florida (ULLZ 5962), Naples, Florida
(ULLZ 5961),Tampa, Florida (ULLZ 5274 to
5276, 5960), Santa Teresa, Florida (ULLZ
3946, 5273), Port St. Joe, Florida (ULLZ
5901); Chandeleur Island, Louisiana (ULLZ
57), Lake Grand Ecaille, Louisiana (ULLZ
2073), California Bay, Louisiana (ULLZ
2075), Breton Island, Louisiana (ULLZ 82),
Grand Isle, Louisiana (ULLZ 3779), Lake
Pelto, Louisiana (ULLZ 2072, 3101, 5277);
Sea Rim State Park, Texas (ULLZ 3889),
Port Aransas, Texas (ULLZ 531, 552, 1564,
2076, 2077, 5282 to 5284, 5958, 5963), Aran-
sas Pass, Texas (ULLZ 1103, 1104, 5285 to
5292), Kline?s Point, Aransas, Texas (ULLZ
921, 2074), Stedman Island, Texas (ULLZ
924, 1766, 1767), Corpus Christi, Texas
(ULLZ 1563, 2071, 5528), Mustang Island,
Texas (ULLZ 1566, 2069), Port Isabel, Texas
(ULLZ 474, 716, 1565), South Padre Island,
Texas (ULLZ 1664, 2067, 5280, 5281, 5293),
U.S.A. Nuevo Torno Grande, Tabasco
(ULLZ 5390); Isla Aguada, Campeche
(ULLZ 1460), Puente Santa Rosal?a, Cam-
peche (ULLZ 4064), Villamar, Campeche
(ULLZ 5395 to 5400), Punta Xen, Campeche
(ULLZ 5391 to 5394), Champot?n, Cam-
peche (ULLZ 2068, 5237, 4056, CNCR 8455),
Campeche, Campeche (ULLZ 2070),
Mexico. Dangriga (ULLZ 5419 to 5423),
Twin Cays (ULLZ 5416 to 5418, 5424, 5425),
Carrie Bow Cay (ULLZ 5426, 5427, 5822),
Belize. Bocas del Toro, Panama (ULLZ
5993, 5995, 5996, 6003, 6093). Bar? (ULLZ
4884), Punta Calixto (ULLZ 5296), Colom-
bia. Cubagua Island (ULLZ 5354, 5969),
Margarita Island (ULLZ 5965, 5967, 5968),
Venezuela. Eastern Pacific: Puertecitos,
Baja California (ULLZ 5329), Santispac,
Muleg?, Baja California Sur, (ULLZ 4137,
5328) Mexico. Potos? (ULLZ 5311 to 5313),
Santa Julia (ULLZ 5314), Estero Aserra-
dores (ULLZ 5310, 5315), Nicaragua. Naos,
Panama City, Panama (ULLZ 5973).
Distribution.?Western Atlantic: Con-
necticut (extralimital); Wilmington, North
Carolina; Sea Island, Georgia; eastern and
western Florida; Lousiana; Texas; U.S.A.;
Tamaulipas, Veracruz, Tabasco, Campeche,
and Quintana Roo, Mexico; Belize; Gulf of
Nicoya, Costa Rica; Bocas del Toro, Col?n,
and Caledonia Bay, Panama; Punta Calixto,
Colombia; Cura?ao; Cubagua and Mar-
garita Islands, Venezuela; Bermuda; Baha-
mas; Cuba; Jamaica; Puerto Rico; Virgin Is-
lands; Bahia, Rio de Janeiro, and Santa
Catarina, Brazil. Eastern Pacific: Sonora
and Baja California in the Gulf of Cali-
fornia, Nayarit, Jalisco, Oaxaca, Mexico;
Puerto el Triunfo and Bah?a de la Uni?n, El
Salvador; Gulf of Fonseca, Honduras; Po-
tos?, Santa Julia, and Estero Aserradores,
Nicaragua; Panama City, Punta Patillo,
Fortified, Toboga, and Flamenco Islands,
Panama; Buenaventura, Colombia; Bah?a
and Punta Santa Elena, Ecuador; Gal?pagos
Islands; Matapalo and Las Vacas, Peru.
Eastern Atlantic: Gibraltar; Senegal to An-
gola; Ascension Island.
Remarks.?Petrolisthes armatus is the most
widespread species in the western Atlantic
and also has a wide range in the eastern
Pacific. It is the only porcellanid species in
Bermuda and it also occurs in the eastern
Atlantic (Africa). It was reported by Smith
(1880) from as far north as Connecticut, an
extralimital record for the western Atlantic
(Haig 1960). In the last decade the species
established in South Carolina, showing a
tendency to extend northward (Knott et al.
2000). The record from Wilmington, North
Carolina, confirms this tendency as well.
Santa Catarina, Brazil, a subtropical area, is
the southern limit of its distribution in the
western Atlantic. Populations of this spe-
cies from different localities show consid-
erable morphological variation. Particu-
larly, the manus of the chelipeds varies
from slender to robust and with spines and
setae along the outer margin to completely
unarmed and hairless. These variations,
however, also occur between specimens of
different sizes from the same locality. As
Haig (1960) reported for Pacific materials,
morphological variations could not be cor-
related with geography. In contrast, based
on molecular data (Rodr?guez et al. in re-
view), three distinct lineages were found,
I. T. RODR?GUEZ ET AL.560
which correspond to defined geographic
regions: the Warm-temperate and Carib-
bean Provinces in the western Atlantic, and
the eastern Pacific. These lineages might in
the future be regarded as distinct species,
provided preliminary molecular evidence
for separations is strongly supported by ad-
ditional molecular and morphological
studies.
Petrolisthes bolivarensis Werding and
Kraus, 2002
Petrolisthes bolivarensis Werding and
Kraus, 2002:1141-1147, figs. 1, 2 (type local-
ity San Mart?n de Pajarales Island, Rosario
Islands, Colombia).
Petrolisthes sp. Werding et al., 2003:79-85,
tables 1, 2.
Material examined.?None.
Distribution.?San Bernardo and Rosario
Islands, Colombia.
Petrolisthes caribensis Werding, 1983
Plate 1, fig. G
Petrolisthes caribensis Werding, 1983a:411,
figs. 1, 4 (type locality Rosario Islands, Co-
lombia).?Hern?ndez, 1999:249, table 2.?
Werding et al., 2003:79-85, tables 1, 2.?
Kraus et al., 2004:85-90.?Rodr?guez et al.,
in review.
Material examined.?Boca Raton (ULLZ
5264), Looe Key Reef (ULLZ 5265, 5266),
American shoal (ULLZ 2852), Florida,
U.S.A. Enmedio Island, Veracruz (ULLZ
5447), Puerto Morelos, Quintana Roo
(CNCR 4895, 5675, 9701), Mexico. Twin
Cays (ULLZ 5428), Carrie Bow Cay (ULLZ
4222, 4317, 4852, 5429), Belize. Bocas del
Toro, Panama (ULLZ 5994, 5997, 6004,
6006, 6091).
Distribution.?Boca Raton and Florida
Keys, Florida, U.S.A.; Veracruz and Quin-
tana Roo, Mexico; Twin and Carrie Bow
Cays, Belize; Bocas del Toro, Panama; Old
Providence and Rosario Islands, and Santa
Marta, Colombia; Venezuela; Puerto Rico.
Remarks.?The first records of this spe-
cies for Mexico and Belize are presented.
Petrolisthes columbiensis Werding, 1983
Petrolisthes columbiensis Werding, 1983a:
413, figs. 2, 5 (type locality Rosario Islands,
Colombia).?Hern?ndez, 1999:249, table
2.?Werding et al., 2003:79-85, tables 1, 2.
Material examined.?None.
Distribution.?Rosario Islands, Colombia;
Cuba.
Petrolisthes dissimulatus Gore, 1983
Petrolisthes cessacii.?Werding, 1982:442
(in part).
Petrolisthes dissimulatus Gore, 1983:94,
figs. 2-4 (type locality St. John, Virgin Is-
lands) .?Werding, 1984:9 , f ig . 3 .?
Hern?ndez, 1999:249, table 2.?Werding et
al., 2003:79-85, tables 1, 2.
Material examined.?None.
Distribution.?Old Providence and Rosa-
rio Islands, and Santa Marta, Colombia;
Cura?ao; Cuba; Puerto Rico; Virgin Islands;
Martinique; Barbados.
Petrolisthes galathinus (Bosc, 1802)
Plate 1, fig. H
Porcellana galathina Bosc, 1802:231-233, pl.
6, fig. 2 (type locality unknown).
Petrolisthes galathinus.?Haig, 1956:22.?
Chace, 1956a:152.?Holthuis, 1959:162.?
Haig, 1960:22, 25, 36-39, 41, 260-261, 386,
387, pl. 19, fig.4, table 6.?Tabb and Man-
ning, 1961:599.?Haig, 1962:175.?Pounds,
1961:28.?Leary, 1964:28.?Bullis and
Thompson, 1965:10.?Williams, 1965:107,
fig. 83.?Co?lho, 1966:56.?Haig, 1966:
352.?Fausto-Filho, 1968:43.?Gore, 1970a:
964.?McCloskey, 1970:30.?Rouse, 1970:
141.?Co?lho, 1971:233.?Co?lho and
Ramos, 1972:174.?Felder, 1973:33-35, pl. 4,
fig. 14.?Gore, 1974:712.?Rickner, 1975b:
164.?Gore and Abele, 1976:21.?Camp et
al., 1977:30.?Werding, 1977:201, fig. 20.?
Werding, 1978b:219.?Young, 1978:176.?
Corredor et al., 1979:32.?Felder and
Chaney, 1979:8, 15, 25, 28-29, fig. 4, tables 1,
2, appendices I, II.?Huni, 1979:21-40, figs.
1-4.?Rodr?guez, 1980:217.?Gore, 1982:
13.?Scelzo, 1982:1133.?Wenner and Read,
1982:187, table 2.?Werding, 1983b:443.?
Werding, 1984:10.?Williams, 1984:243-244,
fig. 178.?Abele and Kim, 1986:413, 420,
421, fig. b.?Raz Guzm?n et al., 1986:349.?
Young, 1986:103, 111.?Ruppert and Fox,
1988:249-250, 404.?Abele and Kim, 1989:
21.?Silva et al., 1989:137, figs. 6, 15.?
PORCELLANIDAE (CRUSTACEA) 561
Williams et al., 1989:35.?Markham et al.,
1990:427.?Mart?nez-Iglesias and Alcolado,
1990:table 26.?Veloso and Melo, 1993:
179.?Arana et al., 1996:217.?Hern?ndez-
Aguilera et al., 1996:50.?Camp et al., 1998:
144.?Garc?a et al., 1998:28-29, table 1.?
Alvarez et al., 1999:9.?Hern?ndez, 1999:
246, 249, tables 1, 2.?Hern?ndez et al.,
1999:27, table 1.?Melo, 1999:252-253, figs.
171, 172.?Stillman and Somero, 2000:200-
208, figs. 1-4.?Mu?oz, 2001:1-53.?
Stillman and Reeb, 2001:236-245, figs. 1, 2,
tables 1-3.?Werding et al., 2001:108-109,
fig. 2, table 3.?L?pez-Greco et al., 2002:40-
46.?Werding and Hiller, 2002:849-850.?
Nizenski, 2003:117.?Werding et al., 2003:
79-85, tables 1, 2.?Hiller et al., 2004:131,
134-135, table 1.?Galicia-Castillo and Her-
na?ndez-Aguilera, 2005:240, 251, fig. 14.?
Rodr?guez et al., in review.
Petrolisthes sexspinosus Gibbes, 1850:
190.?Pounds, 1961:28.?Leary, 1964:28 (as
per Gore and Abele, 1976, this record as-
signable to P. galathinus).
Petrolisthes palathinus.?Werding, 1977:
208 (misspelling).
Petrolisthes galatinus.?Hern?ndez-
Aguilera and Sosa-Hern?ndez, 1982:39-40,
fig. 19. (misspelling).
Not Petrolisthes galathinus.?Haig, 1960:
34, 47, 50.
Material examined.?Atlantic: Florida
Keys, Florida (ULLZ 1986, 1991, 2850, 4524,
4525, 5267 to 5271, 5983); 71?2 Fathom Reef,
Texas (ULLZ 2054, 2055, 2057 to 2064),
U.S.A. Barra del Tordo, Tamaulipas (ULLZ
5820); Topatilla Reef, off Ant?n Lizardo,
Veracruz (ULLZ 5238), Canc?n, Quintana
Roo (ULLZ 5449), Puerto Morelos, Quin-
tana Roo (ULLZ 2056, CNCR 9718, 10937),
Majahual, Quintana Roo (ULLZ 5401),
Mexico. Carrie Bow Cay, Belize (ULLZ
4851, 5430 to 5432). Bocas del Toro, Panama
(ULLZ 6000, 6087, 6090, 6099, 6105, 6109).
Cubagua Island (ULLZ 5355, 5970), Mar-
garita Island (ULLZ 5356), Venezuela. Pa-
cific: Estero Aserradores, Nicaragua (ULLZ
5316).
Distribution.?Western Atlantic: From
Cape Hatteras, North Carolina, South
Carolina, Florida Keys, Texas, U.S.A.;
Tamaulipas, Veracruz, Campeche, Yucata?n,
and Quintana Roo, Mexico; Carrie Bow
Cay, Belize; Bocas del Toro, Col?n, and
Caledonia Bay, Panama; Rosario Islands
and Santa Marta, Colombia; Aruba,
Cura?ao, and Bonaire; Los Roques, Mar-
garita, and Cubagua Islands, Venezuela;
Trinidad; Cuba; Jamaica; Puerto Rico; Vir-
gin Islands; Barbados; Pernambuco, Ala-
g?as, Bahia, Rio de Janeiro, Sta. Catarina,
and Trinidade, Brazil. Eastern Pacific: San
Lucas Island, Costa Rica; Estero Aserra-
dores, Nicaragua; Panama City and Isla To-
baguilla, Panama; La Libertad, Ecuador.
Remarks.?Petrolisthes galathinus has been
treated as a widespread species ranging
from North Carolina, U.S.A., to Santa Ca-
tarina, Brazil, in the western Atlantic (Haig
1956; Williams 1984), and from San Lucas
Island, Costa Rica, to La Libertad, Ecuador,
in the eastern Pacific (Abele and Kim 1989;
Hiller et al. 2004). Various authors recog-
nized the presence of different color forms
within the species. Benedict (1901) de-
scribed two different color forms from
Puerto Rico; Rickner (1975b) noted consid-
erable color variation in specimens from
the east coast of Mexico as did Abele and
Kim (1989) for specimens from the Car-
ibbean and Pacific coast of Panama; and
Williams (1984) cited different color forms
described by other authors. Werding (1977)
was the first to attempt systematic revisions
of this species complex, initially separating
P. rosariensis Werding, 1978, on the basis of
different coloration and presence of a sec-
ond epibranchial spine. Two additional
species were described based on coloration,
epibranchial spination, and the number of
movable spines on the dactylus of the
walking legs: P. caribensis Werding, 1983,
and P. columbiensis Werding, 1983. Two
more variations were recently described: P.
sanmartini Werding and Hiller, 2002, and P.
bolivarensis Werding and Kraus, 2002. This
complex includes species from the eastern
Pacific (P. glasselli) and the Indo-Pacific (P.
bispinosus, P. boscii, P. decacanthus, P. el-
dredgei, and P. moluccensis), accounting for a
total of 12 species (Werding and Hiller, un-
published). More sibling species from the
Caribbean and the Gulf of Mexico remain
to be described. The identity of P. galathinus
continues to be unclear, in part because the
species was superficially described by Bosc
I. T. RODR?GUEZ ET AL.562
(1802). Gibbes (1854) provided a detailed
description and suggested that the original
material might have originated from the
southern Atlantic coast of the United States.
Werding and Hiller (2002) postulated
?? as the typical form of P. galathinus a
morph exhibiting a striped colour pattern,
bearing one epibranchial spine, a supraocu-
lar and an infraocular spine, and three
movable spines on the dactylus of all walk-
ing legs?. However, those characters are
not sufficient to distinguish P. galathinus
from P. bolivarensis and other diverse
morphs that remain undescribed. The dis-
tribution and identity of P. galathinus re-
mains questionable (Hiller et al. 2004). In
another aspect, the reproduction of speci-
mens from Venezuela is currently under
study (I. G. Hern?ndez-?vila et al. unpub-
lished).
Petrolisthes gertrudae Werding, 1996
Petrolisthes gertrudae Werding, 1996:306,
figs 1, 2 (type locality Guadeloupe).?
Werding et al., 2003:79-85, tables 1, 2.
Material examined.?None.
Distribution.?Bonaire and Guadeloupe.
Petrolisthes jugosus Streets, 1872
Plate 1, fig. I
Petrolisthes jugosus Streets, 1872:134 (type
locality St. Martin Island).?Haig, 1962:
180.?Rickner, 1975b:164.?Gore and
Abele, 1976:22.?Werding, 1977:303, fig.
21.?Werding, 1978b:219.?Corredor et al.,
1979:32.?Gore, 1982:14.?Scelzo, 1982:
1134.?Werding, 1982:443.?Werding,
1984:11.?Abele and Kim, 1986:413, 420,
421, fig. a.?Williams et al., 1989:35.?
Markham et al., 1990:427.?Mart?nez-
Iglesias et al., 1993:13.?Hern?ndez-
Aguilera et al., 1996:51.?Camp et al., 1998:
144.?Alvarez et al., 1999:9.?Hern?ndez,
1999:249, table 2.?Werding et al., 2001:108-
109, table 3, fig. 2.?Mag?n et al., 2003.?
Werding et al., 2003:79-85, tables 1, 2.?
Galicia-Castillo and Herna?ndez-Aguilera,
2005:240, 251-252, fig. 15.?Rodr?guez et al.,
in review.
Pisosoma jugosum.?Haig, 1956:16.?
Rodr?guez, 1980:215.
Material examined.?Looe Key Reef,
Florida Keys, U.S.A. (ULLZ 4823). Punta
Delgada, Veracruz (ULLZ 5402, 5452), El
Morro, Veracruz (ULLZ 5239, 5240), La
Mancha, Veracruz (ULLZ 5403, 5404), Es-
condida Beach, Veracruz (ULLZ 5405), En-
medio Island, Veracruz (ULLZ 4850), Can-
c?n, Quintana Roo (ULLZ 4843), Puerto
Morelos, Quintana Roo (ULLZ 5407, 5408),
Majahual, Quintana Roo (ULLZ 5406),
Mexico. Carrie Bow Cay, Belize (ULLZ
5433). Bocas del Toro, Panama (ULLZ 6092,
6100, 6106). Punta Brava, Colombia (ULLZ
4482). Macanao Peninsula, Margarita Is-
land, Venezuela (ULLZ 5357).
Distribution.?The Florida Keys and
Charlotte Harbor, western Florida, U.S.A.;
Veracruz, Campeche, Yucat?n, and Quin-
tana Roo, Mexico; Carrie Bow Cay, Belize;
Bocas del Toro and Caledonia Bay,
Panama; Santa Marta and Punta Brava, Co-
lombia; Cura?ao and Bonaire; La Tortuga,
Margarita, and Cubagua Islands, Venezu-
ela; Tobago; Cuba; Puerto Rico; St. Martin;
Virgin Islands; Barbados.
Remarks.?Morphological differences
were observed between material examined
from Panama and specimens from other lo-
calities. The chelipeds and walking legs are
more densely setose and the carapace has
more rugae on the lateral margins from epi-
branchial to posterior regions, in the Pana-
manian materials.
Petrolisthes magdalenensis Werding, 1978
Petrolisthes sp. II Werding 1977:209, fig. 27.
Petrolisthes magdalenensis Werding, 1978a:
307, fig. 1 (type locality Chengue Bay, Santa
Marta, Colombia).?Gore, 1982:16.?
Scelzo, 1982:1134.?Scelzo, 1983:80.?
Mu?ller and Werding, 1990:257-270, figs.
1 - 7 . ? O s a w a , 1 9 9 5 : 1 6 7 , t a b l e 5 . ?
Hern?ndez, 1999:246, 249, tables 1, 2.?
Werding et al., 2001:108-109, fig. 2, table
3.?Werding et al., 2003:79-85, tables 1, 2.?
Galicia-Castillo and Herna?ndez-Aguilera,
2005:237, 241, 252-253, fig. 16.?Rodr?guez
et al., in review.
Petrolisthes lewisi.?Mag?n, 2001:1-39.
Material examined.?Roat?n, Bay Islands,
Honduras (ULLZ 5442). Macanao Penin-
sula, Margarita Island, Venezuela (ULLZ
5358).
PORCELLANIDAE (CRUSTACEA) 563
Distribution.?Quintana Roo, Mexico;
Roat?n, Bay Islands, Honduras; Portobelo,
Panama; Gulf of Dari?n, Cartagena, and
Santa Marta, Colombia; Margarita Island,
Venezuela; Trinidad and Tobago.
Remarks.?The first record of the species
for Honduras is reported. Morphological
differences between Petrolisthes magdalenen-
sis and P. lewisi are to be published (G.
Hern?ndez et al. unpublished).
Petrolisthes marginatus Stimpson, 1859
Petrolisthes marginatus Stimpson, 1858:
227 (nomen nudum); 1859:74 (type locality
Barbados).?Haig, 1956:26-27 (in part).?
Chace, 1962:620, 622.?Rodr?guez, 1980:
218.?Scelzo, 1982:1134.?Gore, 1982:17.?
Gore, 1983:91, fig. 1.?Werding, 1984:12.?
Veloso and Melo, 1993:180.?Garc?a et al.,
1998:28-29, table 1.?Alvarez et al., 1999:
9.?Melo, 1999:254-255, figs. 173, 174.?
Werding et al., 2003:79-85, tables 1, 2.?
Galicia-Castillo and Herna?ndez-Aguilera,
2005:240, 253-254, fig. 17.?Rodr?guez et al.,
in review.
Petrolisthes cessacii A. Milne-Edwards,
1878:229.?Chace, 1956b:14-17, fig. 4.?
Co?lho, 1971:233.?Co?lho and Ramos,
1972:173.?Gore, 1974:710.?Rickner,
1975b:163.?Werding, 1977:176, 197, 199-
200, fig. 19.?Corredor et al., 1979:32.?
Scelzo, 1982:1133.?Scelzo, 1983:80.?
Hern?ndez, 1999:249, table 2.?Melo, 1999:
250-251, figs. 169, 170.
Not Petrolisthes marginatus.?Haig, 1960:
47-50, pl. 20, fig. 1, table 10.?Carvacho,
1980:250, table 1.
Material examined.?Punta Delgada
(ULLZ 5241), El Morro (ULLZ 5242), La
Mancha (ULLZ 5409, 5410), Escondida
Beach (ULLZ 5411), Montep?o (CNCR
17749), Veracruz, Mexico. El Maguey, Ma-
canao Peninsula, Margarita Island, Venezu-
ela (ULLZ 5359).
Distribution.?Western Atlantic: Vera-
cruz, Mexico; Gulf of San Bl?s, Panama;
Old Providence and Rosario Islands, and
Santa Marta, Colombia; Cura?ao; Los
Roques, Aves, and Margarita Islands, Ven-
ezuela; Trinidad and Tobago; Puerto Rico;
Barbados; Sao Luis and Fernando de No-
ronha, Brazil. Eastern Atlantic: Cape Verde
Islands to Annobon Island, off western Af-
rica.
Remarks.?Petrolisthes haigae Chace, 1962,
is the geminate species of P. marginatus in
the Pacific. The identity of P. marginatus
was reviewed by Gore (1983), who con-
cluded that P. cessacii was its junior syn-
onym. The larval development of P. margi-
natus is currently being described (M.
Pi?ate et al. unpublished).
Petrolisthes politus (Gray, 1831)
Porcellana polita Gray, 1831:14 (type local-
ity not designated).
Petrolisthes politus.?Haig, 1956:21.?
Haig, 1962:178.?Gore, 1974:713, fig. 5.?
Rickner, 1975b:164.?Werding, 1977:203,
fig. 22.?Corredor et al., 1979:32.?
Rodr?guez, 1980:217.?Scelzo, 1982:1134.?
Werding, 1982:444.?Werding, 1984:
12.?Abele and Kim, 1986:413, 420, 421, fig.
d.?Williams et al., 1989:35.?Hern?ndez et
al., 1995:2.?Hern?ndez-Aguilera et al.,
1996:51.?Camp et al., 1998:144.?Garc?a et
al., 1998:28-29, table 1.?Alvarez et al., 1999:
9.?Hern?ndez, 1999:246, 249, tables 1, 2.?
Hern?ndez et al., 2000:143-156, figs. 1-6
tables 1, 2.?Werding et al., 2003:79-85,
tables 1, 2.?Galicia-Castillo and Herna?n-
dez-Aguilera, 2005:241, 254, fig. 18.?
Rodr?guez et al., in review.
Material examined.?Tuxpan, Veracruz
(ULLZ 5243), Enmedio Reef, Veracruz
(ULLZ 5742), Topatilla Reef, off Ant?n Li-
zardo, Veracruz (ULLZ 5244), Canc?n,
Quintana Roo (ULLZ 4847), Chemmuyil,
Quintana Roo (CNCR 9814), Majahual,
Quintana Roo (ULLZ 5412), Mexico. Carrie
Bow Cay, Belize (ULLZ 5434). El Man-
glecito, Cubagua Island, Venezuela (ULLZ
5360).
Distribution.?Florida Keys, U.S.A.; Vera-
cruz and Quintana Roo, Mexico; Carrie
Bow Cay, Belize; Col?n, Panama; Old
Providence and Rosario Islands, and Santa
Marta, Colombia; Aruba, Cura?ao, and Bo-
naire; Los Roques, Aves, and Cubagua Is-
lands, Venezuela; Tobago; Bahamas; Puerto
Rico; Virgin Islands; Antigua; Guadeloupe;
Barbados.
Remarks.?The first record of the species
is reported.
I. T. RODR?GUEZ ET AL.564
Petrolisthes quadratus Benedict, 1901
Petrolisthes quadratus Benedict, 1901:134,
pl. 3, fig. 4 (type locality Ponce, Puerto
Rico).?Haig, 1956:18.?Haig, 1962:179.?
Chace and Hobbs, 1969:121, fig. 32.?
Rickner, 1975b:164.?Werding, 1977:204,
fig. 23.?Corredor et al., 1979:32.?
Rodr?guez, 1980:216.?Scelzo, 1982:1134.?
Werding, 1982:444.?Markham et al., 1990:
427.?Hern?ndez-Aguilera et al., 1996:51.?
Alvarez et al., 1999:9.?Hern?ndez, 1999:
249, table 2.?Werding et al., 2003:79-85,
tables 1, 2.?Galicia-Castillo and Herna?n-
dez-Aguilera, 2005:240, 255, fig. 19.?
Rodr?guez et al., in review.
Material examined.?Looe Key (ULLZ
1987), Pelican Shoals (ULLZ 5272), Florida,
U.S.A. Cozumel Island, Quintana Roo,
Mexico (CNCR 3724). Carrie Bow Cay, Be-
lize (ULLZ 4221). La Tortuga Island, Ven-
ezuela (ULLZ 5361).
Distribution.?Florida Keys, U.S.A.; Vera-
cruz, Campeche, Yucat?n, and Quintana
Roo, Mexico; Carrie Bow Cay, Belize; Cale-
donia Bay, Panama; Old Providence and
Rosario Islands, and Santa Marta, Colom-
bia; Aruba, Cura?ao, and Bonaire; La Tor-
tuga and Cubagua Islands, Venezuela;
Trinidad; Bahamas; Puerto Rico; Virgin Is-
lands; Dominica.
Remarks.?The first records of the species
for U.S.A. and Belize are presented, and its
distribution is extended northward.
Petrolisthes robsonae Glassell, 1945
Petrolisthes robsonae Glassell, 1945:227,
fig. 3 (type locality Miraflores Locks,
Panama Canal, Panama).?Haig, 1960:57,
pl. 18, fig. 2.?Haig, 1968:65.?Gore and
Abele, 1974:567, fig. 3c.?Gore and Abele,
1976:24.?Carvacho, 1980:250, table 1.?
Gore, 1982:19.?Moran, 1984:80.?Abele
and Kim 1989:22.?Hern?ndez-Alvarez,
1995:52-53, pl. 13, fig. 2.?Werding et al.,
2003:79-85, table 1.?Garc?a-Guerrero et al.,
2005:339-349, figs. 1-6, table 1.
Material examined.?None.
Distribution.?Western Atlantic: Gatun
Locks, Panama. Eastern Pacific: Oaxaca,
Mexico; El Salvador; Miraflores Locks,
Panama City, Panama; Guayaquil, Ecua-
dor.
Remarks.?This species is widely distrib-
uted in the eastern Pacific, with occasional
migrations through the Panama Canal up
to Gatun Locks. In the western Atlantic,
Haig (1960) was the first to report a single
specimen collected by Hildebrand in 1935.
Later, Abele and Kim (1989) reported 4 ju-
veniles in the same locality.
Petrolisthes rosariensis Werding, 1978
Petrolisthes sp. I Werding, 1977:208, fig.
26.
Petrolisthes rosariensis Werding, 1978b:214
(type localities Santa Marta and Nen-
guange, Colombia).?Corredor et al., 1979:
33.?Gore, 1982:19.?Werding, 1982:444,
fig. 3 (?-see Remarks below).?Werding,
1983a:410.?Young, 1986:103, 111.?Veloso
and Melo, 1993:180.?Hern?ndez, 1999:249,
table 2.?Melo, 1999:256-257, figs. 175,
176.?Werding et al., 2003:79-85, tables 1,
2.?Rodr?guez et al., in review.
Material examined.?Carrie Bow Cay, Be-
lize (ULLZ 5435, 5445). Bocas del Toro,
Panama (ULLZ 5974).
Distribution.?Carrie Bow Cay, Belize;
Lim?n Bay and Bocas del Toro, Panama;
Rosario Islands and Santa Marta, Colom-
bia; Paraiba to Bahia, Brazil.
Remarks.?The first record of this species
for Belize is presented and its distribution
is extended northward. The taxonomic his-
tory of P. rosariensis is confusing, in large
part due to the publication of the name and
association of this name with a description
well in advance of the date intended by the
author. Because Werding (1978) used the
name P. rosariensis and associated it with
materials listed, illustrated, and briefly di-
agnosed in a previously published account
for ?Petrolisthes sp. I? (Werding 1977), the
name became valid in 1978. That publica-
tion date meets minimum criteria for a pub-
lished description well in advance of the
date that the intended description of this
species (Werding 1982) was actually pub-
lished. Thus, the materials listed in Wer-
ding (1977) become the syntypes for this
species. In acknowledgement of this situa-
tion, Werding (1983a) invoked Article 16 of
PORCELLANIDAE (CRUSTACEA) 565
the ICZN in an attempt to designate as a
lectotype the specimen listed as the holo-
type in Werding (1982); however, this does
not appear to be a justifiable case under the
Code, and the types must thus remain
those presumed syntypes from Santa Marta
and Nenguange, Colombia, listed and de-
scribed in Werding (1977). The ?holotype?
and ?paratypes? listed for P. rosariensis
from the Rosario Islands, Colombia, as des-
ignated in Werding (1982), could be re-
garded as simply additional materials as-
signed to this species, were if not for yet
other complications. The illustration for
Petrolisthes sp. I in Werding (1977:fig. 26)
shows unmistakable presence of a strong
supraocular spine on each side of the cara-
pace, along with a pair of epibranchial
spines. The only North Atlantic species in
which this condition is reported to occur is
P. columbiensis Werding, 1983, yet another
species based on materials from the Rosario
Islands; however, other characters would
appear to exclude it from being confused
with P. rosariensis. Werding (1982:444)
notes specifically in the course of describ-
ing materials as P. rosariensis that there is
?No strong superocular spine?? Unfortu-
nately, in subsequent publications (Wer-
ding (1983b), Werding and Hiller (2002),
Werding and Kraus (2002), and Werding et
al. (2003)) no attempt was made to rectify
this disparity in accounts. Thus, the iden-
tity of P. rosariensis remains somewhat un-
certain. For the present, we must conserva-
tively adhere to the description as
published in Werding (1977) for assign-
ment of our specimens. In so doing, we are
assigning our materials with a strong su-
praocular spine to P. rosariensis. At the
same time, some of the literature records
included above are likely based upon ma-
terials that lack this spine, and may thus
represent a second species. All of the fol-
lowing are possibilities: The illustration in
Werding (1977) or Werding (1982) may
have been in error; the development of the
supraocular spine may be variable; or, an
additional undescribed species may be rep-
resented by materials listed in Werding
(1982) or our materials and perhaps others
cited above. Only study of the original syn-
types is likely to resolve this uncertainty.
Petrolisthes sanmartini Werding and Hiller,
2002
Petrolisthes sanmartini Werding and
Hiller, 2002:849-857, figs. 1-3 (type locality
San Martin Island, Rosario Islands, Colom-
bia).?Werding et al., 2003:79-85, tables 1, 2.
Material examined.?Carrie Bow Cay, Be-
lize (USNM 335750). Bocas del Toro,
Panama (ULLZ 5999).
Distribution.?Carrie Bow Cay, Belize;
Bocas del Toro, Panama; Rosario Islands,
Colombia.
Remarks.?The first records of the species
for Belize and Panama are presented and
the distribution of the species is extended
northward.
Petrolisthes tonsorius Haig, 1960
Petrolisthes tonsorius Haig, 1960:85, pls. 3,
26, fig. 1, table 24 (type locality Berkeley
Cape, Albemarle Island, Gal?pagos Is-
lands).?Haig, 1968:57, 66.?Gore and
Abele, 1976:8, 13, table 3.?Werding, 1977:
205, fig. 24.?Werding, 1978b:220.?Brusca,
1980:263, 265, fig. 17.9.?Carvacho, 1980:
250, table 1.?Gore, 1982:20.?Scelzo, 1982:
1134.?Scelzo, 1983:80.?Pellegrini and
Gamba, 1985:251-267, figs. 1-8.?Lemaitre
and Alvarez-Le?n, 1992:49, table 1.?
Hern?ndez-Alvarez, 1995:54-55, pl. 14, fig.
2.?Osawa, 1995:167, table 5.?Hern?ndez,
1999:246, 249, tables 1, 2.?Hern?ndez et al.,
1999:27, table 1.?Werding et al., 2001:108-
109, fig. 2, table 3.?Werding et al., 2003:
79-85, tables 1, 2.?Rodr?guez et al., in re-
view.
Material examined.?Western Atlantic:
Boca del R?o, Margarita Island, Venezuela
(ULLZ 5362). Eastern Pacific: Puerto Val-
larta, Jalisco, Mexico (ULLZ 5340).
Distribution.?Western Atlantic: Gulf of
Dari?n and Santa Marta, Colombia; Mar-
garita Island and central coast of Venezu-
ela. Eastern Pacific: Baja California Sur, Re-
villagigedo Islands off Colima, Jalisco,
Guerrero, and Oaxaca, Mexico; Port Parker
and Cocos Islands, Costa Rica; Malpelo Is-
land, Colombia; Punta Santa Elena and
Gal?pagos Islands, Ecuador.
Remarks.?Petrolisthes tonsorius was origi-
nally described from Gal?pagos Islands, in
I. T. RODR?GUEZ ET AL.566
the tropical eastern Pacific. It is a species
widely distributed in both tropical coasts of
the Americas.
Petrolisthes tridentatus Stimpson, 1859
Petrolisthes tridentatus Stimpson, 1858:227
(nomen nudum), 1859:75, pl. 1, fig. 4 (type
locality St. Thomas and Barbados).?Haig,
1956:18-19.?Haig, 1960:81-83, pl. 25, fig.
4.?Haig, 1962:179.?Haig, 1968:57, 65.?
Gore, 1971b:485-501, figs. 1-6.?Gore,
1972a:186-208, figs. 36-41, table 12.?Gore
and Abele, 1976:24.?Werding, 1977:208,
fig. 25.?Corredor et al., 1979:33.?
Rodr?guez, 1980:215.?Gore, 1982:20.?
Scelzo, 1982:1134.?Werding, 1982:444.?
Werding, 1984:13.?Lemaitre and Alvarez-
Le?n, 1992:49, Table 1.?Osawa, 1995:167,
table 5.?Hern?ndez, 1999:246, 249, tables
1, 2.?Stillman and Somero, 2000:200-208,
figs. 1-4.?Stillman and Reeb, 2001:236-245,
figs. 1, 2, tables 1-3.?Werding et al., 2003:
79-85, tables 1, 2.?Rodr?guez et al., in re-
view.
Material examined.?Western Atlantic:
Majahual, Quintana Roo, Mexico (ULLZ
5413 to 5415). Falmouth, Jamaica (ULLZ
5441). Boca del R?o, Margarita Island, Ven-
ezuela (ULLZ 5363). Eastern Pacific: Estero
Nagualapa (ULLZ 5324), Estero Aserra-
dores (ULLZ 5306, 5307), Nicaragua. Naos
Island, Panama (ULLZ 5364, 5975).
Distribution.?Western Atlantic: Quin-
tana Roo, Mexico; Caledonia Bay, Panama;
Old Providence and Rosario Islands, and
Santa Marta, Colombia; Margarita and
Cubagua Islands, Venezuela; Trinidad; Ba-
hamas; Cuba; Jamaica; St. Thomas; Anti-
gua; Barbados. Eastern Pacific: Salinas Bay
and Port Parker, Costa Rica; San Juan del
Sur, Nicaragua; Honda Bay, Tobaguilla Is-
land, Perlas Islands, and Guayabo Chi-
quito, Panama; Lim?n Bay, Colombia;
Santa Elena Bay and Pun? Island, Ecuador.
Remarks.?The first records of the species
for Mexico and Jamaica are presented.
Petrolisthes tridentatus comprises a species
complex and cryptic species remain to be
described (Rodr?guez et al. in review).
Genus Pisidia Leach, 1820
Pisidia Leach, 1820:53 (type species Pi-
sidia linnaeana Leach, 1820, designation by
Haig, 1960).?Haig, 1960:207-209.?
Rodr?guez et al., in review.
Porcellanides Czerniavsky, 1884:109 (type
species Porcellanides kriczagini Czerniavsky,
1884, by original designation).
Streptochirus Stimpson, 1907:188 (type
species Porcellana serratifrons Stimpson,
1858, designation by Haig, 1960).
Remarks.?Pisidia brasiliensis Haig in Rod-
rigues da Costa (1968) from Brazil was the
first species of the genus reported in the
western Atlantic. It has also been found in
the southern Caribbean Sea (Werding 1977
and this report). A second species from Bra-
zil, P. melloleitaoi Rodrigues da Costa, 1968,
could not be confirmed as valid (Veloso
and Melo 1993). P. magdalenensis (Glassell,
1936) is the only representative of the genus
in the eastern Pacific.
Pisidia brasiliensis Haig (in Rodrigues da
Costa, 1968)
Pisidia brasiliensis Haig (in Rodrigues da
Costa, 1968):406-R (type localities Capanea
and Sao Sebastiao, Sao Paulo, Bra-
zil).?Co?lho, 1971:233.?Werding, 1977:
211, fig. 28.?Scelzo, 1982:1132.?Scelzo,
1983:80.?Veloso and Melo, 1993:180.?
Hern?ndez, Bola?os et al., 1996:21.?
Hern?ndez, 1999:246, 249, tables 1,
2.?Melo, 1999:258-259, figs. 177, 178.?
Werding et al., 2003:79-85, tables 1, 2.
Pisidia sp.?: Co?lho and Ramos, 1972:175.
Material examined.?Macanao Peninsula,
Margarita Island, Venezuela (ULLZ 5365,
5971, 5972).
Distribution .?Cartagena, Ci?naga
Grande de Santa Marta, and Santa Marta,
Colombia; Margarita Island, Venezuela;
Par? to Pernambuco, Brazil.
Remarks.?The species was briefly de-
scribed without an illustration; a detailed
description of the species is to be published
(C. Lira et al., unpublished). The popula-
tion structure, reproductive aspects, and
the larval development of the species are
currently under study (G. Hern?ndez et al.,
unpublished).
Genus Polyonyx Stimpson, 1858
Polyonyx Stimpson, 1858:229 (type spe-
cies Porcellana macrocheles Gibbes, 1854, by
PORCELLANIDAE (CRUSTACEA) 567
original designation).?Haig, 1960:232-
233.?Rodr?guez et al., in review.
Polyonx.?Menzel, 1971:79 (misspelling).
Remarks.?Only one species of Polyonyx
occurs in the western Atlantic, P. gibbesi
Haig, 1956; three other species of the genus
occur in the eastern Pacific: P. confinis Haig,
1960, P. nitidus Lockington, 1878, and P.
quadriungulatus Glassell, 1935. The species
in this genus live as commensals in the
tubes of Chaetopterus (Annelida: Poly-
chaeta) (Haig 1960; Werding 1983b).
Polyonyx gibbesi Haig, 1956
Porcellana macrocheles Gibbes, 1850:191
(type locality South Carolina, U.S.A.).
Polyonyx gibbesi Haig, 1956:28.?Haig,
1960:238-239.?Gray, 1961:353-359.?
Williams, 1965:113, fig. 90.?Co?lho, 1966:
63.?Haig, 1966:356.?Gore, 1968:111-129,
figs. 1-8.?Rouse, 1970:141.?Dudley and
Judy, 1971:3-6, 9, tables 1-3, 5.?Gosner,
1971:539, fig. 21.53A.?Co?lho and Ramos,
1972:174.?Van Engel and Sandifer, 1972:
158.?Felder, 1973:33-36, pl. 4, fig. 10.?
Sandifer, 1973:245.?Craig, 1974:235-243.?
Gore, 1974:713.?Caine, 1975:283, fig. 3.?
Rickner, 1975a:313-314.?Young, 1978:
176.?Felder and Chaney, 1979:8, 15, 25,
appendix I.?Scelzo, 1982:1132.?Maris,
1983:237-246, figs. 1-7.?Truesdale and An-
dryszak, 1983:43, table 2.?Williams, 1984:
244, fig. 179.?Abele and Kim, 1986:410,
424, 425, fig. d.?Ruppert and Fox, 1988:
204-205, figs. 201, 250, 404.?Britton and
Morton, 1989: 193-194, fig 7.10 F.?
Williams et al., 1989:35.?Mart?nez-Iglesias
et al., 1993:13.?Veloso and Melo, 1993:
181.?Camp et al., 1998:144.?Hern?ndez,
1999:246, 249, tables 1, 2.?Melo, 1999:260-
261, figs. 179, 180.?Nizenski, 2003:117.?
Werding et al., 2003:79-85, tables 1, 2.?
Rodr?guez et al., in review.
Polyonx macrocheles.?Menzel, 1971:79
(misspelling).
Polyonyx gibbesii.?Rodrigues da Costa,
1964, p. 325 (misspelling).
Polyonix gibbesi.?Silva et al., 1989:137,
figs. 7, 12 (misspelling).
Material examined.?Fort Pierce, eastern
Florida (ULLZ 5254 to 5259), Turkey Pt.,
Florida (ULLZ 3559), Destin, Florida (ULLZ
1989), U.S.A.
Distribution.?Massachusetts, Rhode Is-
land, North Carolina, South Carolina, east-
ern and western Florida, Texas, U.S.A.;
Caledonia Bay, Panama; Punta Papuy, Ven-
ezuela; Trinidad; Cuba; Puerto Rico; Rio de
Janeiro to southern Brazil; La Paloma, Uru-
guay.
Remarks.?The distribution of this species
is mostly antitropical; it is sporadically
found in tropical waters. In this study,
slight morphological differences in the
carapace were observed between speci-
mens from both sides of the Florida Penin-
sula.
Genus Porcellana Lamarck, 1801
Porcellana Lamarck, 1801:153 (type spe-
cies Cancer platy-cheles Pennant, 1777, by
monopyty).?Haig, 1960:196-197.?Haig,
1978:706-714.?Rodr?guez et al., in review.
Platycheles Billberg, 1820:134 (type spe-
cies Cancer platy-cheles Pennant, 1777, by
tautonymy).
Enostea Gistel, 1848:159, 196 (substitute
name for Porcellana Lamarck, 1801; taking
the same type species, Cancer platy-cheles
Pennant, 1777).
Remarks.?Porcellana sayana (Leach,
1820), P. sigsbeiana A. Milne-Edwards, 1880,
and P. stimpsoni A. Milne-Edwards, 1880,
were included in Haig?s (1956) report for
the western North Atlantic. Porcellana pai-
vacarvalhoi Rodrigues da Costa, 1968, was
later incorporated in the western Atlantic
revision of the genus (Haig 1978). Subse-
quently, P. stimpsoni was considered a jun-
ior synonym of P. sayana in Lemaitre and
Campos (2000), and P. paivacarbalhoi was
considered a junior synonym of P. platyche-
les by Veloso and Melo (1993). Only one
male represented the latter species in the
western Atlantic; this species is better
known from the eastern Atlantic (Haig
1978; Veloso and Melo 1993). The report of
only one specimen of P. platycheles in the
western Atlantic raises questions about its
recognition as a component of the porcel-
lanid fauna of the region. Additional rec-
ords are desirable in order to confirm the
establishment of the species in this region.
With the recent description of P. lillyae Le-
maitre and Campos, 2000, the genus con-
tains three species in the western Atlantic.
I. T. RODR?GUEZ ET AL.568
Porcellana lillyae Lemaitre and Campos,
2000
Porcellana lillyae Lemaitre and Campos,
2000:259-265, figs. 1, 2 a-c, 3 a-c, g, h, 4 a-c
(type locality Gulf of Morrosquillo, Colom-
bia).?Werding et al., 2003:79-85, tables 1, 2.
Material examined.?None.
Distribution.?Reported only from the
Caribbean coast of Colombia between Gulf
of Morrosquillo and San Bernardo Islands.
Porcellana sayana (Leach, 1820)
Pisidia sayana Leach, 1820:54 (type locali-
ties Georgia and Florida).
Porcellana sayana.?Haig, 1956:31-33.?
Chace, 1956a:152.?Holthuis, 1959:161.?
Haig, 1962:186.?Co?lho, 1964:255.?Leary,
1964:27-28.?Bullis and Thompson, 1965:
10.?Williams, 1965:110, fig. 87.?Co?lho,
1966:62.?Haig, 1966:354.?Gore, 1970a:
963.?Rouse, 1970:141.?Co?lho, 1971:
233.?Menzel, 1971:79.?Co?lho and Ra-
mos, 1972:175.?Felder, 1973:34-36, pl. 4,
fig. 15.?Gore, 1974:715.?Fotheringham
and Brunenmeister, 1975:61, 63, 167, fig.
3.23.?Rickner, 1975b:162.?Voss, 1976:92-
93, fig.?Werding, 1977:212, fig. 29.?Haig,
1978:707.?Young, 1978:176.?Corredor et
al., 1979:33.?Felder and Chaney, 1979:25,
fig. 4, appendix I.?Rodr?guez, 1980:218.?
Hern?ndez-Aguilera and Sosa-Hern?ndez,
1982:41-42, fig. 20.?Scelzo, 1982:1132.?
Wenner and Read, 1982:186, 194, 195, tables
2, 5.?Werding, 1982:446.?Maris, 1983:
237-246, figs. 1-7.?Werding, 1984:13.?
Williams, 1984:245, fig. 180.?Abele and
Kim, 1986:413, 422, 423, fig. b.?Ruppert
and Fox, 1988:250, 404.?Britton and Mor-
ton, 1989: 245, 247, 258, fig. 10-1 G1.?Silva
et al., 1989:138, figs. 9, 13.?Williams et al.,
1989:35, pl. 1, third appendix page.?
Markham et al., 1990:427.?Human, 1992:
164-165, middle color plate.?Veloso and
Melo, 1993:182.?Hern?ndez-Aguilera et
al., 1996:53.?Tunnell and Alvarado, 1996:
136.?Camp et al., 1998:144.?Hern?ndez et
al., 1998:101-118, tables 1-3, figs. 1-7.?
Alvarez et al., 1999:9.?Hern?ndez, 1999:
246, 249, tables 1, 2.?Hern?ndez et al.,
1999:27, table 1.?Melo, 1999:266-267, figs.
183, 184.?Lemaitre and Campos, 2000:264,
figs. 2f, 3f .?Nizenski, 2003:117.?Werding
et al., 2003:79-85, tables 1, 2.?Galicia-
Castillo and Herna?ndez-Aguilera, 2005:237,
240, 255-256, fig. 20.?Rodr?guez et al., in
review.
Porcellana sayana?.?Gore, 1983:22.
Porcellana stimpsoni A. Milne-Edwards,
1880:35.?Haig, 1956:33.?Haig, 1978:
707.?Abele and Kim, 1986:413, 422, 423,
fig. a.?Williams et al., 1989:35.?Camp et
al., 1998:144.?Lemaitre and Campos, 2000:
264.
Material examined.?Bethel Shoal, off Fort
Pierce, eastern Florida (ULLZ 5260), off
Sanibel Island, Florida (ULLZ 1908), off
Destin, Florida (ULLZ 2044), Florida (ULLZ
267); off Mississippi, (ULLZ 1101, 1102,
1701, 1809, 2047); North Gulf of Mexico
(ULLZ 1350, 1351); Grand Isle, Louisiana
(ULLZ 3100), Eugene Island, Louisiana
(ULLZ 392), offshore Louisiana (ULLZ
4574); off Port Aransas, Texas (ULLZ 2046),
North Padre Island, Texas (ULLZ 2049), UT
Longhorn, Texas (ULLZ 1515), 7 1?2 Fathom
Reef, Texas (ULLZ 2045, 2050, 2052, 2053),
U.S.A. Off Champot?n, Campeche, Mexico
(ULLZ 2051). Twin Cays, Belize (ULLZ
5436 to 5438). Guayacanecito, Macanao
Peninsula, Margarita Island, Venezuela
(ULLZ 5366).
Distribution.?North Carolina, South
Carolina, Georgia, eastern and western
Florida, Mississippi, Louisiana, and Texas,
U.S.A.; Tamaulipas, Veracruz, Campeche,
and Yucat?n, and Quintana Roo, Mexico;
Twin Cays, Belize; Caledonia Bay, Panama;
La Guajira, De La Vela Cape, and Santa
Marta, Colombia; Los Roques, Margarita,
Cubagua, and Coche Islands, Venezuela;
Bahamas; Cuba; Jamaica; Puerto Rico; Vir-
gin Islands; Antigua; Barbados; Guyanas;
southward to Rio Grande do Sul, Brazil.
Remarks.?The first record of this species
for Belize is presented. As noted by Le-
maitre and Campos (2000), the status of
Porcellana stimpsoni is reviewed by Werding
(Werding in preparation).
Porcellana sigsbeiana A. Milne-Edwards, 1880
Porcellana sigsbeiana A. Milne-Edwards,
1880:35 (type localities off delta of Missis-
sippi, north of Yucat?n, and Flannegan Pas-
PORCELLANIDAE (CRUSTACEA) 569
sage, Virgin Islands).?Haig, 1956:33.?
Bullis and Thompson, 1965:10.?Williams,
1965:111, fig. 88.?Dawson, 1966:177, table
1.?Gore, 1970a:964.?Pequegnat and
Chace, 1970: 162-164, 166, 168, tables 5-2,
5-4.?Gore, 1971c:344-355, figs. 1-6.?
Gosner, 1971:539.?Gore, 1972a:122-156,
figs. 24-29, tables 8, 9.?Van Engel and San-
difer, 1972:158.?Felder, 1973:34-36, pl. 4,
fig.16.?Gore, 1974:716.?Haig, 1978:707.?
Young, 1978:176.?Co?lho et al., 1980:39.?
Scelzo, 1982:1132.?Wenner and Read,
1982:187, 194, tables 2, 5.?Maris, 1983:
239.?Truesdale and Andryszak, 1983:43,
49, table 2.?Williams, 1984:246, fig. 181.?
Abele and Kim, 1986:413, 422, 423, fig. c.?
Williams et al., 1989:35.?Veloso and Melo,
1993:182.?Hern?ndez-Aguilera et al., 1996:
53.?Camp et al., 1998:144.?Alvarez et al.,
1999:9.?Hern?ndez, 1999:246, 249, tables 1,
2.?Melo, 1999:268?269, figs. 185, 186.?
Lemaitre and Campos, 2000:259, 264, figs.
2d, e, 3d, 4d.?Nizenski, 2003:117.?
Werding et al., 2003:79-85, tables 1, 2.?
Lalana et al., 2004:6, table 1.?Castillo and
Herna?ndez-Aguilera, 2005:240, 256-257, fig.
21.?Rodr?guez et al., in review.
Material examined.?Straits of Florida
(ULLZ 4481), Gulf of Mexico (ULLZ 736,
1352), off Mississippi (ULLZ 1663, 4837), off
Louisiana (ULLZ 1990, 5448, 5455, 5902),
U.S.A.
Distribution.?Massachusetts, North
Carolina, Florida, Mississippi, Louisiana,
northern Gulf of Mexico, U.S.A.; Tamauli-
pas, Veracruz, Tabasco, Campeche, and
Yucat?n, Mexico; Honduras; Colombia;
Venezuela; Trinidad; Cuba; Virgin Islands;
Suriname; Par? and Maranhao, Brazil.
DISCUSSION
The porcellanid fauna of the western At-
lantic numbers 48 valid species plus two
undescribed species herein reported (Neo-
pisosoma sp. and Pachycheles sp). Of the
western Atlantic species described since
Haig (1956), 17 have been subsequently
recognized as valid (Megalobrachium
mortenseni, Neopisosoma neglectum, N. orien-
tale, Pachycheles chubutensis, P. cristobalensis,
P. susanae, Petrolisthes bolivarensis, P. cari-
bensis, P. columbiensis, P. dissimulatus, P. ger-
trudae, P. magdalenensis, P. rosariensis, P.
sanmartini, P. tonsorius, Pisidia brasiliensis,
and Porcellana lillyae), six species have been
determined to be junior synonyms (Mega-
lobrachium walteri, Pachycheles haigae,
Petrolisthes cessacii, P. serratus, Porcellana
paivacarvalhoi, and P. stimpsoni), two have
been considered invalid (Petrolisthes costai
and Pisidia melloleitaoi), and one (Porcellana
platycheles) has been excluded because only
one specimen has been reported question-
ably for the western Atlantic. Similarly, 11
genera are now known for the region, three
of these (Neopisosoma, Clastotoechus, and
Parapetrolisthes) were recognized after
Haig?s revisions (Haig 1956, 1960, 1962). Pi-
sidia was resurrected and Porcellanopsis was
combined with Megalobrachium (Haig
1960). The monospecific genus Madarateu-
chus, established by Harvey (1999) to re-
ceive Clastotoechus vanderhorsti, is not found
justified.
Regarding the porcellanids from Florida
and the Gulf of Mexico, this study shows
that the family is well represented in the
literature for the region. Only Petrolisthes
quadratus is herein reported for the first
time in Florida and its range is extended
northward to the U.S.A. However, previ-
ously unreported intra-specific morpho-
logical variation was observed among
some species. Several morphotypes of P.
galathinus occur in Florida, Texas, Veracruz,
and other localities of this study. These
likely correspond to undescribed species
currently confused under a P. galathinus-
complex of species (Werding 1982, 1983a;
Werding and Hiller 2002; Werding and
Kraus 2002).
Another case of intra-specific morpho-
logical variability is that of Polyonyx gibbesi.
Although few specimens were available for
examination, differences in the shape of the
carapace were observed between samples
from the two sides of the Florida Peninsula.
Different populations or cryptic species
within P. gibbesi might be recognized, as are
known among populations of other trans-
Floridian decapods of the genera Sesarma,
Uca, Callichirus, and Pagurus (see Felder
and Staton 1994; Staton and Felder 1995;
Young et al. 2002). Preliminary molecular
evidence of divergence was observed in
I. T. RODR?GUEZ ET AL.570
trans-Floridian specimens of Megalobra-
chium soriatum, which suggests that this
taxon may contain cryptic species (Rod-
r?guez et al. in review). In this case how-
ever, no morphological variation could be
correlated to geography.
Examination of materials from Quintana
Roo, on the Caribbean coast of Mexico, Be-
lize, and Honduras resulted in the discov-
ery of an undescribed species of the genus
Neopisosoma and the range extension of
three species northward: Megalobrachium
mortenseni, Petrolisthes rosariensis, and P.
sanmartini. Three species are recorded for
the first time in Mexican waters: Petrolisthes
caribensis, P. tridentatus, and the unde-
scribed species of Neopisosoma. Seven spe-
cies are new for Belize: Pachycheles riisei,
Petrolisthes caribensis, P. politus, P. quadratus,
P. rosariensis, P. sanmartini, and Porcellana
sayana. The two species from Honduras,
Megalobrachium mortenseni and Petrolisthes
magdalenensis, were not previously known
in the area. These findings confirm that the
porcellanid fauna in this region remains
poorly known.
Porcellanids from the southern Carib-
bean are fairly well known (Werding
1996; Werding et al. 2003). Even so, in this
study an undescribed species of the genus
Pachycheles is reported from Panama, and
Petrolisthes sanmartini is reported for the
first time from that country. In the Antilles,
Petrolisthes tridentatus is recorded for the
first time from Jamaica.
Fourty-eight valid species plus two un-
described species are currently known for
the western Atlantic. Particularly variable
are Petrolisthes galathinus, P. armatus, and P.
tridentatus. Petrolisthes galathinus is now
known to consist of a complex of at least six
species in the southern Caribbean Sea: P.
bolivarensis, P. caribensis, P. columbiensis, P.
rosariensis, P. sanmartini, and P. galathinus
itself (see Werding and Hiller 2002). The
latter continues to group a series of mor-
phologically and genetically variable forms
still in need of more detailed revision (Still-
man and Reeb 2001; Rodr?guez et al. in re-
view).
Petrolisthes armatus shows considerable
morphological variation throughout its
wide range (Chace 1956b; Haig 1956, 1960).
Evidence from the morphology of the lar-
vae suggests that the populations are diver-
gent. The larvae of Atlantic and Pacific
specimens described by Gore (1970b,
1972b) differ in several important features.
Similarly, zoeal and megalopal stages of P.
armatus from Venezuelan islands differed
in several characteristics from those from
Bermuda, Florida, the Gulf of Mexico, and
the eastern Pacific (Graterol 1996). Speci-
mens of P. armatus from the Gulf of Cali-
fornia and from each coast of Panama were
included in a molecular study by Stillman
and Reeb (2001), where specimens from the
two Pacific localities grouped more closely
together than with Atlantic specimens, to
the exclusion of other species of the genus.
An additional third population distributed
in the northern Gulf of Mexico and eastern
coast of the U.S.A. was suggested in pre-
liminary molecular studies (Rodr?guez et
al. in review). Although molecular data
have shown that populations of P. armatus
present differences throughout their
ranges, and thus might be regarded as dis-
tinct species, morphological examinations
of extensive series from the eastern Pacific
(Haig 1960) and the western Atlantic (this
study) concluded that variations in mor-
phology did not consistently correlate with
geography.
The third species-complex, Petrolisthes
tridentatus, is also amphi-American and has
been subject to morphological and genetic
comparative studies that showed differ-
ences between specimens from both sides
of the Isthmus of Panama (Gore 1971b;
Stillman and Reeb 2001). Materials exam-
ined in this study from the northern Carib-
bean Sea and from Venezuela were mor-
phologically distinct from those from
Panama. This distinction was supported in
preliminary molecular analyses (Rodr?guez
et al. in review). These findings coupled
with additional morphological and mo-
lecular analyses, are likely to result in the
description of new species.
Biogeographic considerations
The porcellanid fauna in the western At-
lantic is for the most part Caribbean (Wer-
ding et al. 2003). Of the recorded species in
PORCELLANIDAE (CRUSTACEA) 571
T
A
B
L
E
2.
D
is
tr
ib
u
ti
on
of
p
or
ce
la
in
cr
ab
s
by
zo
og
eo
gr
ap
h
ic
p
ro
v
in
ce
s
in
th
e
w
es
te
rn
A
tl
an
ti
c.
P
ro
v
in
ce
s
m
od
if
ie
d
af
te
r
B
ri
gg
s
(1
97
4)
,W
il
li
am
s
(1
98
4)
,a
n
d
B
os
ch
i
(2
00
0)
as
fo
ll
ow
s:
V
ir
gi
n
ia
n
,
fr
om
C
ap
e
C
od
to
C
ap
e
H
at
te
ra
s;
N
W
ar
m
-t
em
p
er
at
e,
ea
st
er
n
U
.S
.A
.
fr
om
C
ap
e
H
at
te
ra
s
to
C
ap
e
C
an
av
er
al
(=
C
ar
ol
in
ia
n
);
G
u
lf
of
M
ex
ic
o
W
ar
m
-t
em
p
er
at
e,
fr
om
C
ap
e
R
om
an
o
to
C
ap
e
R
oj
o,
M
ex
ic
o;
C
ar
ib
be
an
,f
ro
m
C
ap
e
R
oj
o
to
O
ri
n
oc
o
D
el
ta
;W
es
tI
n
d
ia
n
,t
h
e
C
ar
ib
be
an
Is
la
n
d
s;
B
ra
zi
li
an
,f
ro
m
th
e
O
ri
n
oc
o
D
el
ta
to
C
ap
e
Fr
io
;
an
d
A
rg
en
ti
n
ia
n
,
fr
om
C
ap
e
Fr
io
to
43
/
4?
S.
T
h
e
oc
cu
rr
en
ce
of
p
or
ce
ll
an
id
s
in
th
e
ea
st
er
n
P
ac
if
ic
is
m
ar
ke
d
in
th
e
fi
n
al
co
lu
m
n
.
Sp
ec
ie
s
V
ir
gi
n
ia
n
N
W
ar
m
-
te
m
p
er
at
e
G
u
lf
of
M
ex
ic
o
W
ar
m
-
te
m
p
er
at
e
C
ar
ib
be
an
W
es
t
In
d
ia
n
B
ra
zi
li
an
A
rg
en
ti
n
ia
n
E
as
te
rn
P
ac
if
ic
C
la
st
ot
oe
ch
us
no
do
su
s
(S
tr
ee
ts
,
18
72
)
?
?
C
la
st
ot
oe
ch
us
va
nd
er
ho
rs
ti
(S
ch
m
it
t,
19
24
)
?
?
E
uc
er
am
us
pr
ae
lo
ng
us
St
im
p
so
n
,
18
60
?
?
?
M
eg
al
ob
ra
ch
iu
m
m
or
te
ns
en
i
H
ai
g,
19
62
?
?
M
eg
al
ob
ra
ch
iu
m
po
ey
i
(G
u
?r
in
,
18
55
)
?
?
?
M
eg
al
ob
ra
ch
iu
m
ro
se
um
(R
at
h
bu
n
,
19
00
)
?
?
?
M
eg
al
ob
ra
ch
iu
m
so
ri
at
um
(S
ay
,
18
18
)
?
?
?
?
?
M
in
yo
ce
ru
s
an
gu
st
us
(D
an
a,
18
52
)
?
?
?
N
eo
pi
so
so
m
a
an
gu
st
ifr
on
s
(B
en
ed
ic
t,
19
01
)
?
?
N
eo
pi
so
so
m
a
cu
ra
ca
oe
ns
e
(S
ch
m
it
t,
19
24
)
?
?
N
eo
pi
so
so
m
a
ne
gl
ec
tu
m
W
er
d
in
g,
19
86
?
?
N
eo
pi
so
so
m
a
or
ie
nt
al
e
W
er
d
in
g,
19
86
?
N
eo
pi
so
so
m
a
sp
.
?
P
ac
hy
ch
el
es
ac
kl
ei
an
us
A
.
M
il
n
e-
E
d
w
ar
d
s,
18
80
?
?
?
P
ac
hy
ch
el
es
ch
ac
ei
H
ai
g,
19
56
?
?
?
P
ac
hy
ch
el
es
ch
ub
ut
en
si
s
B
os
ch
i,
19
63
?
P
ac
hy
ch
el
es
cr
is
to
ba
le
ns
is
G
or
e,
19
70
?
P
ac
hy
ch
el
es
gr
ee
le
yi
(R
at
h
bu
n
,
19
00
)
?
P
ac
hy
ch
el
es
la
ev
id
ac
ty
lu
s
O
rt
m
an
n
,
18
92
?
?
P
ac
hy
ch
el
es
m
on
ili
fe
r
(D
an
a,
18
52
)
?
?
?
P
ac
hy
ch
el
es
pi
lo
su
s
(H
.
M
il
n
e
E
d
w
ar
d
s,
18
37
)
?
?
?
P
ac
hy
ch
el
es
ri
is
ei
(S
ti
m
p
so
n
,
18
59
)
?
?
?
P
ac
hy
ch
el
es
ru
gi
m
an
us
A
.
M
il
n
e-
E
d
w
ar
d
s,
18
80
?
?
?
P
ac
hy
ch
el
es
se
rr
at
us
(B
en
ed
ic
t,
19
01
)
?
?
P
ac
hy
ch
el
es
su
sa
na
e
G
or
e
an
d
A
be
le
,
19
73
?
?
P
ac
hy
ch
el
es
sp
.
?
P
ar
ap
et
ro
lis
th
es
to
rt
ug
en
si
s
(G
la
ss
el
l,
19
45
)
?
?
P
et
ro
lis
th
es
am
oe
nu
s
(G
u
?r
in
,
18
55
)
?
?
?
P
et
ro
lis
th
es
ar
m
at
us
(G
ib
be
s,
18
50
)
?
?
?
?
?
?
?
?
P
et
ro
lis
th
es
bo
liv
ar
en
si
s
W
er
d
in
g
an
d
K
ra
u
s,
20
02
?
I. T. RODR?GUEZ ET AL.572
T
A
B
L
E
2.
C
on
ti
n
u
ed
.
Sp
ec
ie
s
V
ir
gi
n
ia
n
N
W
ar
m
-
te
m
p
er
at
e
G
u
lf
of
M
ex
ic
o
W
ar
m
-
te
m
p
er
at
e
C
ar
ib
be
an
W
es
t
In
d
ia
n
B
ra
zi
li
an
A
rg
en
ti
n
ia
n
E
as
te
rn
P
ac
if
ic
P
et
ro
lis
th
es
ca
ri
be
ns
is
W
er
d
in
g,
19
83
?
?
P
et
ro
lis
th
es
co
lu
m
bi
en
si
s
W
er
d
in
g,
19
83
?
?
P
et
ro
lis
th
es
di
ss
im
ul
at
us
G
or
e,
19
83
?
?
P
et
ro
lis
th
es
ga
la
th
in
us
(B
os
c,
18
02
)
?
?
?
?
?
?
?
P
et
ro
lis
th
es
ge
rt
ru
da
e
W
er
d
in
g,
19
96
?
?
P
et
ro
lis
th
es
ju
go
su
s
St
re
et
s,
18
72
?
?
P
et
ro
lis
th
es
m
ag
da
le
ne
ns
is
W
er
d
in
g,
19
78
?
P
et
ro
lis
th
es
m
ar
gi
na
tu
s
St
im
p
so
n
,
18
59
?
?
?
P
et
ro
lis
th
es
po
lit
us
(G
ra
y,
18
31
)
?
?
P
et
ro
lis
th
es
qu
ad
ra
tu
s
B
en
ed
ic
t,
19
01
?
?
P
et
ro
lis
th
es
ro
bs
on
ae
G
la
ss
el
l,
19
45
?
?
P
et
ro
lis
th
es
ro
sa
ri
en
si
s
W
er
d
in
g,
19
82
?
?
P
et
ro
lis
th
es
sa
nm
ar
ti
ni
W
er
d
in
g
an
d
H
il
le
r,
20
02
?
P
et
ro
lis
th
es
to
ns
or
iu
s
H
ai
g,
19
60
?
?
P
et
ro
lis
th
es
tr
id
en
ta
tu
s
St
im
p
so
n
,
18
59
?
?
?
P
is
id
ia
br
as
ili
en
si
s
H
ai
g,
19
68
?
?
P
ol
yo
ny
x
gi
bb
es
i
H
ai
g,
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PORCELLANIDAE (CRUSTACEA) 573
this study, only four have not been found
in the Caribbean Province. Euceramus
praelongus is entirely a northern temperate
species, Pachycheles chubutensis is exclu-
sively distributed in the southern temper-
ate zone, P. laevidactylus lives in this zone
but also reaches tropical Brazil, and P. gree-
leyi is endemic to tropical Brazil (Table 2).
Zoogeographic affinities of porcelain crabs
were discussed by Werding et al. (2003),
who noted that the fauna from the West
Indian Province of Briggs (1974) was dis-
tinguished from that of the southern Car-
ibbean by having a lower species number
(34 versus 40 species). In this study we
found that all species in the West Indian
Province (32 species) were also present in
the Caribbean Province (46 species) (Table
2). Therefore only 14 species in the Car-
ibbean were not found in the West Indies.
Because some of these species were re-
cently described, it is probable that they too
will be found there eventually. Some of the
Caribbean species clearly have a wider dis-
tribution than previously thought (e.g.,
Minyocerus angustus, Neopisosoma orientale,
Petrolisthes magdalenensis, P. rosariensis, P.
sanmartini, and Pisidia brasiliensis). We sug-
gest that the West Indian Province should
be considered an indistinguishable compo-
nent of the Caribbean Province regarding
the porcellanid fauna.
Knowledge of the Porcellanidae in the
western Atlantic has been improved with
the study of larval development and the
use of molecular data, in addition to tradi-
tional studies of adult morphology. Even
so, detailed studies focused on particular
species and genera are required to clarify
taxonomic status, phylogenetic relation-
ships, and biogeographic patterns. Further
explorations in many parts of Central
America, the Antilles, and the Guyanas are
essential to better understanding porcel-
lanid composition and distribution in the
region.
Acknowledgments.?We are grateful to the
many individuals who assisted in collect-
ing the porcellanids of the ULLZ collection
and especially thank Rafael Lemaitre,
Emilio Garc?a, Suzanne Fredericq, Rafael
Robles, Christoph Schubart, Jos? Cuesta,
Fernando Mantelatto, Fred Gurgel, Joel
Stake, Brian Mahon, Antonio Baeza, Carlos
Lira, Juan Bola?os, Antonio Ord??ez, and
Julieta Flores. Fernando Alvarez and Jos?
Luis Villalobos gracefully facilitated access
to materials of the CNCR and Rafael Le-
maitre those of the USNM. The manuscript
benefited greatly from comments offered
by Raymond Bauer, Joseph Neigel, Rafael
Lemaitre, Fernando Alvarez, and Joel Mar-
tin. This study was supported by U.S. Na-
tional Science Foundation grant no. DEB-
0315995, U.S. Department of Energy grant
no. DE-FG02-97ER12220, and U.S. Depart-
ment of the Interior (Program USGS/AID-
Nicaragua) cooperative agreement award
00CRAG0009 to D. L. Felder at the Univer-
sity of Louisiana at Lafayette, and awards
from the Consejo de Investigaci?n de la
Universidad de Oriente to G. Hern?ndez in
Venezuela.
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