Contributions to Zoology, 69 (4) 223-250 (2000)
SPB Academic Publishing bv, The Hague
Re-evaluation of the Cancridae Latreille, 1802 (Decapoda: Brachyura) including three new genera and three new species
Carrie+E. Schweitzer & Rodney+M. Feldmann
Department of Geology, Kent State University, Kent, Ohio 44242 U.S.A. E-mail: [email protected]. edu and [email protected]
Keywords: Decapoda, Brachyura, Cancridae, Tertiary, paleobiogeography, Tethys
Abstract Metacarcinus A. Milne Edwards, 1862 235
Metacarcinus goederti new species 236
Notocarcinus new genus 239 New fossils referable to the Cancridae Latreille, 1802 extend
Notocarcinus sulcatus new species 240 the known into the middle Eocene stratigraphic range of the family Platepistoma Rathbun, 1906 241 and the into South America. Each within geographicrange genus Romaleon Gistl, 1848 242 the family has been reevaluated within the context ofthe new Lobocarcininae 1930 243 material. of characters for each cancrid Subfamily Beurlen, A suite diagnostic ge- Lobocarcinus Reuss, 1867 243 nus makes it possible to assign both extant and fossil speci- Muller, 1979 244 and the two cancrid the Cancrinae Miocyclus mens to genera subfamilies, Tasadia Muller in Janssen and Miiller, 1984 244 Latreille, 1802, and Lobocarcininae Beurlen, 1930,based solely
Discussion 245 dorsal is upon carapace morphology. Cheliped morphology useful
246 in the but is less useful Acknowledgements assigning genera to family significantly
References 246 at the subfamily and generic level. Each ofthe four subgenera
Appendix A 249 sensu Nations (1975), Cancer Linnaeus, 1758, Glebocarcinus
B 249 Nations, 1975, Metacarcinus A. Milne Edwards, 1862, and Appendix
Addi- Romaleon Gistl, 1848,are elevated to full generic status.
three and three new accommodate tionally, new genera species
the new, as well as some previously described taxa, and include Introduction and Anatolikos new genus, Anisospinos berglundi new genus
and and species, and Notocarcinus sulcatus new genus species
several combinations. of and The Cancridae Latreille, 1802, has heretofore con- new Recognition new genera
several the sisted of known from the fossil reassignment of species within Cancrinae indicates two extant genera
that that subfamily have arisen in the southern hemisphere, lato may record, Cancer sensu and Platepistoma Rath-
contrary to the previous interpretation of the subfamily as a several fossil bun, 1906, and exclusively genera North Pacific The primarily or Tethyan group. Lobocarcininae including Lobocarcinus Reuss, 1867; Microdium was primarily a Tethyan group. Reuss, 1867; Miocyclus, Muller, 1979; Parapirimela
Van Straelen, 1937; and Tasadia Muller in Janssen
and 1984. Cancer lato subdivided Contents Muller, sensu was into four subgenera by Nations (1975) including
Roma- Introduction 223 Cancer, Glebocarcinus, Metacarcinus, and
Systematic 224 leon. All of these and de- Paleontology genera subgenera were Cancridae Family Latreille, 1802 224 fined the and primarily upon number, size, shape Subfamily Cancrinae,Latreille, 1802 226 of the frontal spines; the number, size, ornament, Cancer sensustricto Linnaeus, 1758 227 and of the anterolateral the ornament Anatolikos shape spines; new genus 228 of the the new posterolateral margin; shape, develop- Anisospinos genus 229 and Anisospinos berglundi new species 231 ment, and ornament of carapace regions, the
1975 234 Glebocarcinus Nations, shape and ornament of the chelae.
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The discovery of new fossil material fromAlaska Systematic Paleontology and Washington, USA, and southern Argentina
the Within this Order 1802 requires a re-evaluation of family. Decapoda Latreille, work, the four subgenera referred to Cancer sensu Infraorder Brachyura Latreille, 1802 lato are elevated to full generic status. Examination Section Heterotremata Guinot, 1977 of all species referred to the Cancridae necessitates Family Cancridae Latreille, 1802 the transfer of several species to other genera and results in recognition of three new genera to accom- Diagnosis. - Carapace wider than long; anterior modate new material as well as some previously margin with four to six spines, with odd or even described fossil and extant species (Appendix A). number usually diagnostic of subfamilies; antero-
New genera include Anatolikos, Anisospinos, and lateral margin with numerous spines, usually more
Notocarcinus. NeitherParapirimela nor Branchio- than eight; posterolateral margin entire, rimmed lambrus verifiable members of the Cancridae from indis- are or spined; carapace regions ranging
discussed below. the tinct as Additionally, morphology to extremely well-defined; carapace ornament of the chelae has been found to be useful in assigning ranging from smooth to coarsely granular to spined; taxa to the family based upon the overall shape chelae typically with keels, spines or granules on and ornamentation. However, chelae are signifi- manus and fingers. Sternum narrow, triangular in
less useful in be- sternites 3-5 often fused in males. cantly assigning taxa to genera, shape; “Anten-
of the in Antennal cause variability chela morphology. nules folded lengthwise. flagella present,
The evolutionary and biogeographic history of short, more or less hairy. Third maxillipcds usu- the be within the endostome” 1930, group must reinterpreted context ally overlapping (Rathbun, p. of the new fossil material. The discovery of Noto- 176). “Carapace broadly oval or hexagonal. Last
sulcatus and from of for carcinus new genus species pair legs not adapted swimming” (Williams, middle Eocene rocks of constitutes the Argentina 1984, p. 351). earliest known occurrence of the Cancrinae and
of the Can- - also the first known fossil occurrence Remarks. This diagnosis (and all the diagnoses
America. cridae in South Anisospinos berglundi to follow) focus on characters that are typically
other Eocene is the only occurrence known, re- preserved in fossils and that are therefore most ported from late Eocene rocks of the western useful to paleontologists. The main feature unit-
It is that the Cancrinae arose the is a of Washington. possible ing family high degree carapace orna- in the southern hemisphere and subsequently dis- mentation, especially the frontal, anterolateral, and persed northward to the North Pacific and North posterolateral spines and/or rims. Additionally,
Atlantic where it is well-estab- members of the mani of the first Oceans, currently family possess lished. The Cancrinae was previously believed to pereiopods that are equal or subequal and orna- have had a primarily North Pacific (Nations, 1975, mented with smooth, granular, or spined keels. A
all 1979; Carvacho, 1989) or Tethyan (Newman, 1991) key to genera herein included within the family distribution. Most of the to the characteristics genera assigned (Appendix B; Fig. I), employs these
Cancrinae in this Anatolikos new Muller the report, including to distinguish taxa. (1984) placed genus genus,Anisospinos new genus, Cancer sensu stricto, Microdium within the Cancridae, and suggested ,
Platepistoma and Romaleon, apparently originated that it be closely allied to or with , may synonymous in the North Pacific based known region upon Cancer s. l. However, the holotypc for the sole occurrences. of the M. has been lost Assignment European genera species, nodulosum, (Muller, 1984),
and Tasadia the Lobocarcininae confirmation of the status of that is im- Miocyclus to sup- so genus ports the primarily Tethyan distribution of that possible. According to Muller (1984, p. 77), it is subfamily. clear that the taxon does belong within the
Cancridae, but because of the lost type, the genus
is not included in the key.
Van Straelen (1937) and Glaessner (1969) placed
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Fig. I. to the and of the Cancridae the of each The Illustrated key subfamilies genera showing geologic range genus. branching structure of the key does not reflect phylogeny; each node represents a couplet in the dichotomous key (Appendix B).
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Parapirimela within the Cancridae. The sole spe- Remarks. - The Cancrinae, until now, has been a
founded of few cies, P. angolensis van Straelen, 1937, was small subfamily comprised only a genera,
of Cancer l. and Para- upon a fragment dorsal carapace and cheliped. s. Platepistoma as well as
described The anterolateral margin was as possess- pirimela, Microdium, and Branchiolambrus, which
four the last ing anterolateral spines, of which is have been discussed above. This report introduces the a front with five three the in the bifid; spines including poorly new genera to subfamily; addition,
inner orbital and a that developed spines; carapace four of Cancer s. l. Cancer s. Gle- subgenera , s.,
wider than None of these characteristics is long. bocarcinus, Metacarcinus, and Romaleon, are
the The can be seen on illustrations of this taxon. herein elevated to generic status, bringing the
is described as housed in a private col- numberof included and holotype genera to eight. Diagnoses lection and therefore, be lost. Thus, it is im- may, included species for each genus are provided be-
to confirm ofthis taxon within possible placement low. Genera within the subfamily are distinguished of the family. The development carapace regions based the of the frontand frontal upon nature spines; and the of the chela that elongate fingers suggest the shape, size, and number of anterolateral spines; this taxon be more readily allied with the may the nature of the ornamentation of the posterolat- Xanthidae. eral margin; and the development and ornamenta- altus Branchiolambrus Rathbun, 1908, was re- tion of carapace regions. ferred to the based upon the dorsal Parthenopidae Nations introduced the use of four (1975, p. 22) front described cancrid- carapace, but the was as subgenera based upon dorsal carapace features for like (Rathbun, 1908). Glaessner (1969) placed the species of the genus Cancer, believing the
Branchiolambrus within the Cancridae. We con- of the be characters carapace to more conserva- with is cur Rathbun (1908) that this genus most tive evolutionarily than cheliped characters. The with closely allied the parthenopids based upon distinct differences in the nature of the front, the of the the inflated the triangular shape carapace, anterolateral spines, the posterolateral margins, and branchial regions, the trifid front, and the small, the development and ornamentation of dorsal cara- circular orbits. distinct elevation pace are sufficiently to justify
of the subgenera to generic status. In the present Subfamily Cancrinae Latreille, 1802 Cancer work, species assigned to s. l. were placed
into based characteristics of the dor- genera upon Included genera.— Anatolikos new genus; Aniso- sal This has resulted in carapace only. some reas- Cancer spinos new genus; sensu stricto Linnaeus, signments of taxa compared with those of Nations 1758; Glebocarcinus Nations, 1975; Metacarcinus (1975). Milne Notocarcinus A. Edwards, 1862; new genus; Even though Nations (1975) primarily used dorsal Platepistoma Rathbun, 1906; Romaleon Gistl, 1848. define the carapace morphology to four subgenera,
he employed the morphology of the claws and Diagnosis. - Carapace wider than long; front with fingers to erect several new species and to assign five spines including inner orbital spines, axial spine them to subgenera. Important features of the chelae be lower level than other orbits with may at spines; include the number of keels on the outer surface two fissures; anterolateral margin with numerous of the the ornamentation of the the manus; keels; spines, spines fissured or separated to bases; pos- of the of the the spinosity upper margin manus; terolateral margin entire, rimmed, or with one or spinosity ofthe uppermargin of the movable finger; two small spines; posterior margin entire or rimmed; and the number, size, and color of the denticles on be distinct or in- carapace regions variable, may the occlusal surfaces of the fixed and movable fin- distinct, may be ornamented with granules or tu-
However, the subgenera as Nations (1975) bercles be chelae with several keels gers. or may smooth; defined them contain chelae of extremely variable on mani, keels may be smooth, granular, or spined; morphology, indicating that the morphology of the mani ofchelae ranging from short and high to long insufficient chelipeds is to permit confident ge- and slender; fingers of chelae ranging from short neric placement of the species formerly referred and high to long and slender.
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Cancer l. the to s. Furthermore, generic reassign- Cancer sensu stricto Linneaus, 1758 ments made herein based upon dorsal carapace
also result Cancer 625. morphology in groups with variable Linnaeus, 1758, p,
91. For within the Alpheus Weber, 1795, p. cheliped morphology. example, genus Pagurus Berthold, 1827,p. 255, (nonPagurus Fabricius, 1775). Metacarcinus, the mani either resemble those of Trichocera De Haan, 1833, p. 4, 16, ( non Trichocera Meigen, Romaleon or those of Cancer s.s.; however, taxa 1803).
to Metacarcinus based are readily assignable upon H, Milne 412, Platycarcinus Edwards, 1834,p. the the nature of dorsal carapace. Therefore, claw Trichocarcinus Miers, 1879, p. 34, (substituted for Trichocera).
Cancer morphology is not necessarily useful in assigning (Cancer) Nations, 1975, p. 23; Karasawa, 1990, p. 11, pi. 1, figs. 2a-d, 3a-d, 8, 16a-b; Sakumoto, Karasawa, and taxa to genera within the Cancrinae. Some genera
Takayasu, 1992, p. 448, pi. 61, figs. 3a-c; Karasawa, 1993, p. do have chelae that be less for appear to variable; 51, pi. 10, figs. 1-2, 7a-b; Karasawa, 1997, p. 47, pi. 10, figs. 1, those of Glebocarcinus, and example, Platepistoma, 4-5, 7a-b.
Anisospinos new genus may in fact be diagnostic.
Those taxa that have been described from Cancer only Type Species. — pagurus Linnaeus, 1758. claw morphology are herein referred to Cancer s.l.
The of taxa known only from cheliped - validity Diagnosis. Carapace ovate, length 58 to 66 per-
fragments is to determine because they maximum impossible cent carapace width. Front weakly pro- may or may not be synonymous with species known jecting beyond orbits; with five spines including from dorsal material carapace only. Taxa assigned inner orbital spines, inner three spines closely Cancer l. include C. allisoni to s. Nations, 1975; spaced; axial spine at lower level than other fron-
C. C. chaneyi Nations, 1975; coosensis Nations, tal spines, frontal region axially sulcate, sulcus
1975; C. durhami Nations, 1975; C. garthi Na- extending posteriorly from position of medial frontal
1975; C. marri Nations, 1975; C. Fronto-orbitalwidth 22 29 tions, styriacus spine. from to percent
Bittner, 1883; and C. Nations, 1975. maximum yanceyi carapace width; orbits shallow, small,
Cancerites molassicus Quenstedt, 1867 is known with two fissures, directed forward. Anterolateral
from of and is removed only a portion a finger, margin very tightly curved posteriorly; nine ante- from with Collins and Cancer s. l., concurring rolateral spines, separated by fissures, typically
Fraaye (1991). truncated distally but occasionally produced into Of the referred the eight genera to Cancrinae, sharp spines posteriorly, margins of spines entire,
two, and Notocarcinus n. Anisospinos n. gen. gen. granular or serrate. Posterolateral margin weakly are known from the fossil record, and exclusively concave, rimmed, typically with one spine. Poste-
members of these two document extension genera rior margin rimmed, convex. Regions usually in- of the of the into the Eocene. range subfamily distinct, best developed axially; smooth or finely
Notocarcinus sulcatus n. sp. is known from the granular. middle Eocene of and Anisospinos Argentina Manus of chelipeds coarsely granular or smooth, berglundi n. has been collected from late Eocene sp. short; usually with three or four granular ridges rocks of the Each Olympic Peninsula, Washington. on outer surface; upper margin keeled, sometimes
of the other Glebocarcinus, has a genera, except with small sharp or blunt spines; lower margin fossil record into the Miocene. Verifi- extending keeled; distal margin typically extending at approxi-
able occurrences of Glebocarcinus are known only 90 mately degree angle to upper margin. Movable from the Recent. surface finger short; upper granular or smooth,
occlusal surface with large, blunt denticles. Fixed
with central finger short, keel or smooth, occlusal
surface with large, blunt denticles.
Remarks. - of Species Cancer s. s. are united in
that possessing anterolateral spines are deeply fis-
sured, a weakly produced front, and one posterolat-
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The of Anatolikos eral spine. chela members of the genus is new genus
typically short with short fingers and granular or
smooth keels.
referred the Cancer Cancer plebejus was to subgenus (Glebocarcinus) Nations, 1975, p. 23, 44, fig. 30.9;
Metacarcinus by Nations (1975). It is here placed Karasawa, 1990, p. 7, pi. 1, fig. 10; Karasawa, 1993, p. 50, pi. 10, fig. 3; Karasawa, 1997,pi. 11, fig. 7. within Cancer based its of s. s. upon possession Cancer 41.4. (Cancer) Nations, 1975, p. 23, 44, fig. 6, 41.3, fissured spines and a weakly produced front with
small, forward-directed orbits. Cancer borealis was Type Species. - Cancer japonicus Ortmann, 1893 placed within Metacarcinus by Nations (1975); it by original designation. based is here placed within Cancer s. s. upon pos-
session of fissured anterolateral spines and a weakly Diagnosis. - Carapace wider than long; front pro- produced front. duced beyond orbits, with five coalesced spines Packard (1900) reported Cancer proavitus from including inner-orbital spines, separated by fissures; Miocene deposits of the east coast ofNorth America. anterolateral margin with 10-12 spines, spines linear of tubercles the axial It possesses a row on separated by fissures, third and fourth, fifth and regions and tubercles on the hepatic regions, or- sixth, seventh and eighth, and ninth and tenth spines namentation not seen in cancrids. Furthermore, the paired. that is material exhibits an epibranchial ridge or-
namented with several tubercles. These features
Description. - Carapace wider than long, maxi- that the be referrable to the suggest specimen may mum length 70 to 80 percent maximum carapace Portunidae, and in fact, the morphology of the ster- width, carapace widest at position of penultimate nites is typical of portunid taxa. Cancer proavitus or last anterolateral spine, located about three- is therefore removedfrom the Cancridae and placed the distance the the quarters posteriorly on carapace; within the Portunidae; unfortunately, poor pres- regions moderately well-defined, ervation of the front and the anterolateral margins carapace espe- cially axial, protogastric, and metabranchial regions; docs not permit further definition.
record Cancer carapace surface Front The fossil for s. l. was reported granular. projected beyond straight, with five spines including to extend into the Miocene by Glaessner (1969) orbits, nearly
Nations inner-orbital frontal lobate, and (1975). The oldest verifiable occur- spines, spines separated
of Cancer fissures. Fronto-orbital width from 31 to 46 rence s. s., based upon dorsal carapace by
morphology, is Cancer fujinaensis from Miocene percent maximum carapace width; orbits with thick-
rocks of Japan (Sakumoto, Karasawa, and Takayasu, ened rim and two fissures. Anterolateral margin with 1992), and Karasawa (1990) described Cancer convex, most convex posteriorly; 10-12 spines,
tomowoi from early to early middle Miocene rocks separated by fissures, triangular at tips; outer-or-
of Japan based on fingers and a portion of thlp ster- bital spine small, not well developed; second spine
num. North American fossil occurrences include singular, sharp; third and fourth, fifth and sixth,
C. fissus, from the Pliocene of California and C. seventh and eighth, and ninth and tenth spines productus from Pliocene and Pleistocene rocks of paired; eleventh and twelfth spines, if present, sin-
C. Alaska to California. Reports of productus are gular and sharp. Posterolateral margin nearly straight
based claw The entire with only upon fragments. genus ap- or concave, rimmed, or one spine.
pears to have evolved in the North Pacific Ocean, Posterior margin nearly straight, rimmed. Regions perhaps during the Miocene, and subsequently moderately well defined, especially axial, proto-
North dispersed throughout the Pacific and Atlan- gastric, and metabranchial regions; surface of cara- tic Oceans by the Pliocene or Pleistocene. Recent pace granular. of Cancer inhabit the Northeast and South- species Manus of cheliped short; outer surface with three east Pacific Ocean and the North Atlantic Ocean, and scattered granular ridges tubercles; upper the the south- suggesting dispersal across equator to margin with three short spines; lower margin keeled. ern hemisphere and to the Atlantic Ocean through Fixed finger short, medially keeled; large, blunt
the Straits of Panama. denticles on occlusal surface. Movable finger short,
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with blunt small, denticles on occlusal granular, Anisospinos new genus surface.
Cancer 15, Imaizumi, 1962,p. 239, pi. 40, fig.
- Cancer and Goedert, 1992, Etymology. The generic name is taken from the (Romaleon)Berglund p. 4; Karasawa,
1990, p. 6. Greek, “anatolikos,” eastern, recognizing that the
genus appears to be endemic to Japan.
Type species. - Cancer (Romaleon) wahkiakumensis
Berglund and Goedert, 1992, by original designa- Occurrence. - The new genus is currently known tion. only from Japan. The two Recent species, A. japo-
nicus and A. tumifrons have been reported from Diagnosis. - Carapace with five evenly-spaced waters surrounding Japan (Nations. 1975), and A. frontal spines including inner-orbital spines; or- itoigawai is known from the early to early middle bits broad, deeply excavated, with well-developed Miocene of southwestern Japan (Karasawa, 1990; rim; anterolateralmargin with eight spines including 1993). outer-orbital spine, spines separated to bases; third
and fourth, fifth and sixth, and seventh and eighth Remarks. - Both Anatolikos japonicus and A. anterolateral spines paired. Posterolateral margin were referred tumifrons originally to the genus entire, rim weak if dis- Cancer. present; carapace regions Subsequently, Nations (1975) placed C. tinct, granular or ornamented with tubercles. Manus japonicus in the subgenus Cancer (Cancer) and with two to four keels on outer surface and sharp C. tumifrons within Cancer (Glebocarcinus) . Ka- spines on upper margin of manus and movable rasawa (1990) placed C. itoigawai within Cancer finger. (Glebocarcinus) . It is clear that these three species
form related a group within the Cancrinae in pos-
Description. - Carapace hexagonal, wider than long; sessing a nearly straight, produced front composed moderately vaulted longitudinally and transversely. of five coalesced spines separated by fissures. No Front slightly projected beyond orbits, with five other cancrid exhibits frontal of this genus spines inner spines including orbital spines, spines evenly nature. Additionally, the shape and arrangement spaced, medial spine sharp, lateral extend- of spines anterolateral spines in the new genus are unique ing as ridge onto protogastric region, front about within the subfamily. The anterolateral spines are 30 percent maximum carapace width. Orbits di- fissures which be closed separated by may or slightly rected weakly anterolaterally, broad, deeply exca- and open, the orbits exhibit a thickened, broad rim, vated; with wide, well-developed rim; two orbital differentiating Anatolikos from all other genera of fissures; fronto-orbital width from 40-55 the ranging subfamily. Anisospinos new genus, to be de- percent maximum carapace width. Anterolateral scribed below, also has a thickened orbital rim; margin with eight spines including outer-orbital however, the long, sharp, separated, and forward spine; spines separated to bases, granular, often curving spines of Anisospinos immediately distin- curving anteriorly; outer-orbital spine sharp; sec- guish members of that from Anatolikos. genus ond spine long, sharp; third and fourth, fifth and Anatolikos have evolved in appears to Japan sixth spines, seventh and eighth spines paired; each during the early Miocene and has subsequently anterolateral spine of different size and shape from remained endemic The modern to that region. spe- other. Posterolateral any margins entire, very weak cies A. tumifrons is especially similar to the fossil rim if present. Posterior margin convex. Carapace species A. differs from these itoigawai ; A. japonicus regions distinct; granular or ornamented with tu- two species in possessing twelve anterolateral spines bercles. instead of ten and an overall wider carapace. Evo- Chelae equal; manus long, ornamented with two lution within remained the group has apparently to four keels on outer surface; keel on lower mar- relatively conservative, resulting in a small, nar- of manus keel gin serrate; on upper margin with defined rowly genus. several sharp or blunt spines. Fixed finger with
grooves parallel to upper and lower margins, oc-
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clusal surface with denticles of varying sizes. Upper maleon, but differs from it in several aspects. In
margin of movable finger with three distally di- Romaleon, the middle three frontal spines are
rected spines; occlusal surface with several blunt closely spaced, while in Anisospinos, they are more
denticles of varying size. evenly spaced. The lateral frontal spines ofAnisos-
extend the in pinos onto protogastric region a long,
- that is shorter and much Etymology. The generic name is derived from well-developed ridge; ridge
the Greek “anisos,” unequal, and “spinos,” spine, less well-developed in members of Romaleon.
the nature of the anterolateral Members of describing unequal Anisospinos possess deeply excavated,
feature of the wide orbits that have a rim. spines, a diagnostic genus. broad, well-developed
Species of Romaleon have shallower, narrower
- All Occurrence. Anisospinos is known from the late orbits that lack a rim. species of Romaleon have
Eocene Floko River Formation, western Olympic nine unpaired anterolateral spines; in Anisospinos,
Peninsula, Washington, USA, and the Miocene there are eight anterolateral spines, the last six of
Astoria Formation, southwestern Washington, USA which occur in pairs. The posterolateral margin of
(Berglund and Goedert, 1992). One species is known species of Romaleon is ornamented either with a
in from the middle Miocene Odose Formation Ja- beaded rim or a rim plus one posterolateral spine;
has and weak rim if pan (Karasawa, 1990). Anisospinos no spines a very
present at all. The carapace regions of members of
Remarks. - Cancer wahkiakumensis was originally Romaleon are smooth or finely granular, while those
referred to Cancer (Romaleon) by Berglund and of Anisospinos are granular or ornamented with
C. wahkiaku- Goedert (1992) who noted that (R.) large tubercles. Finally, the manus of the chelipeds
mensis differed from all other species of the sub- in Romaleon is ornamented with well-developed
genus Romaleon in the shape of the carapace, ar- keels composed of spines or large granules, whereas
rangement of carapace regions and in anteriorly in Anisospinos, those keels are much more poorly
directed anterolateral spines of differing sizes and developed. Because of these numerous differences,
shapes. Additionally, they noted that C. (R.) wahkia- Anisospinos is easily distinguished from Romaleon.
kumensis possessed only eight anterolateral spines Anisospinos is known from late Eocene and
while other species of Cancer (Romaleon) have Miocene rocks of Washington. Anisospinos odosen-
those observed that nine spines. Finally, authors sis occurs in middle Miocene rocks of Japan. This
C. wahkiakumenis lacks that the evolved in the northeastern (R.) posterolateral spines suggests genus
or which all other of Romaleon Pacific the and rims, species pos- Ocean during Eocene subsequently The sess. discovery of a new species from the Hoko dispersed to Japan via surface ocean currents or
River Formation prompts description of a new along the Aleutian Island Arc. Additional fossil
genus, Anisospinos, which includes C. wahkiaku- occurrences of Anisospinos along the North Pacific
rim will be confirm mensis, C. odosensis, and a new species to be de- necessary to this dispersal
scribed below. pattern.
is from all Anisospinos distinguishable other The Japanese species Anisospinos odosensis was
of Cancridae based its Cancer Imaizumi genera the upon possession originally referred to by (1962)
of eight anterolateral spines that are separated to and was later referred to Cancer (Romaleon) by
in No their bases, the last six of which occur pairs. Karasawa (1990, 1993, 1997). This species is re-
other cancrid exhibits such anterolateral ferrable based its genus to Anisospinos upon possession
ornamentation. the orbits of Anisos- of anterolateral that are Additionally, paired, sharp spines sepa- pinos are deeply excavated, broad, and broadly rated to their bases. Additionally, each anterolateral
suite of orbital characters rimmed, a not exhibited spine is different in size from any other spine. The
other cancrid orbits of C. odosensis by any genus. Finally, Anisospinos are broad, deeply excavated,
differs from all of the in and rimmed. Cancer odosensis genera lacking postero- bifissured, possesses
lateral unrimmed spines or rims. an entire posterolateral margin that is
is Ro- front. Anisospinos most similar to the genus and a produced All of these characters are
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the C. odosensis Anterolateral with diagnostic of new genus; therefore, margin weakly convex, eight is referred to Anisospinos. Karasawa (1993, 1997) spines including outer orbital spine; posterior
he referred First illustrated a specimen (MFM83202) that margins of spines sometimes granular. spine to Cancer sanbonsugii. Examination of that speci- (outer orbital spine) sharp, narrow, directed for- men and comparison to the holotype of Cancer ward. Second anterolateral spine broadly triangu-
directed odosensis indicates that MFM83202 is a member lar, forward, anterior margin concave, of C. based of convex. Third and fourth odosensis, upon possession paired, posterior margin weakly
third sharp and well-developed anterolateral spines, anterolateral spines paired; spine narrow,
from second overall carapace shape, regional shape and develop- attenuated, separated spine by deep, ment, and orbital size and shape. narrow reentrant, directed forward, anterior mar-
gin weakly concave, posterior margin weakly con-
vex; fourth anterolateral spine broadly triangular,
directed from third Anisospinos berglundi new species weakly anterolaterally, separated
(Figs. 2-3) spine by shallow notch. Fifth and sixth anterolat- eral spines paired; fifth spine small, triangular,
directed Diagnosis. - Carapace broadly hexagonal, wider sharp, weakly anterolaterally, separated sixth than long, widest at position of last anterolateral from fourth spine by v-shaped notch; spine
directed antero- spine; frontal margin with five evenly spaced spines broadly triangular, sharp, strongly lower including inner-orbital spines; orbits deeply exca- laterally, uppermargin nearly straight, margin from fifth shallow vated, wide, with broad rim; anterolateral margin convex, separated spine by broad, notch. Seventh and seventh with eight spines including outer-orbital spine; third eighth spines paired; and fourth, fifth and sixth, and seventh and eighth spine narrow, short, sharp, anterior margin weakly
concave, margin weakly directed anterolateral spines paired; posterolateral margin posterior convex,
on anterolaterally; eighth spine longest, atten- entire; chelae equal; mani of chelae with keels strongly uated, directed from seventh tipper and lower margins and outer surface. laterally, separated
spine by u-shaped notch. Short, shallow groove
from between second - onto region Description. Carapace hexagonal, wider than long, extending hepatic
and third anterolateral shallow L/W = of last spines; longer 0.74; carapace widest at position groove
onto from between fourth anterolateral spine, located about two-thirds dis- extending hepatic region
and fifth anterolateral spines; longer, shallow groove tance posteriorly on carapace; carapace regions from between sixth and moderately to weakly inflated, defined by shallow extending onto carapace seventh anterolateral surface with and spines. grooves; carapace sparse granules Posterolateral margins sinuous, concave anteri- scattered larger tubercles; carapace moderately
and convex vaulted transversely and longitudinally. orly posteriorly, entire; posterior margin about 32 maximum Front with five spines including inner orbital weakly convex, percent cara- width. spines; axial spine small, blunt-tipped; spines on pace Protogastric region triangular in shape, moder- either side of axial spine broadestof frontal spines, inflated, ornamentedwith scattered tubercles, separated from axial spine by shallow, u-shaped ately directed with longitudinal ridge parallel and adjacent to notch; innerorbital spines narrow, sharp, mesogastric extending from two frontal forward, separated from frontal spines by deeper, process,
spines on either side of axial spine aboutone-third v-shaped notch; front about 27 percent maximum
distance on inner carapace width. posteriorly protogastric region;
of region weakly convex, outer Fronto-orbital width about 55 percent maximum margin protogastric
convex. narrow carapace width; orbits broad, circular, directed margin weakly Mesogastric region broadened lateral forward, with wide rim; two orbital fissures, inner anteriorly, posteriorly, margins
distal convex. Urogastric orbital fissure at about midwidth of orbit, outer concave, margin weakly inflated, anterior lateral orbital fissure located just proximal to outer or- region margin concave,
margins concave, posterior margin weakly convex. bital spine.
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2. and Anterior view of USNM 507767. Dorsal of USNM Fig. Anisospinos berglundi n. gen. sp. 1, holotype, 2, carapace holotype,
507767. 3, Dorsal carapace of paratype, USNM 507768.
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and lower grooves parallel to upper margins.
Movable finger highest proximally and narrowed
distally, height to length ratio about 0.39; upper
with three first shortest margin convex, spines, spine
and blunt, second and third spines long, narrow
and sharp, all three spines directed distally; oc-
clusal surface concave, with several variably sized
blunt denticles.
Carpus of cheliped slightly longer than high,
globular; outer surface convex, ornamented with
a few tubercles; upper margin with sharp spine at
distal corner and smaller, sharp spine just proximal
remainder of to long spine, upper margin nearly
distal with blunt straight; margin convex, projection
at mid-height; lower margin convex; proximal Fig. 3. Line drawing of Anisospinos berglundi showing the with position and orientation ofmeasurements taken. margin nearly straight, sharp triangular spine
well known. at upper corner; merus not
Remainder of venter and not known. Cardiac region rounded-triangle in shape, apex appendages
directed posteriorly, two large swellings anteriorly,
Measurements. - Measurements taken on one roundedswelling posteriorly. Intestinal region (in mm)
of in flattened, not well differentiated. specimens Anisospinos berglundi are given
Hepatic region weakly inflated, ornamented with Tables 1 and 2. Position and orientation of measure-
taken in 3. scattered tubercles. Epigastric region inflated cen- ments are illustrated Fig.
trally and flattened laterally, ornamentedwith scat-
tered tubercles. Mesobranchial region inflated, Etymology. - The trivial name honors Ross E.
ornamented with scattered tubercles. Metabranchial Berglund, Bainbridge Island, WA, who has made
region flattened, ornamented with several large a significant contribution to the study of the Can-
tubercles, with semi-lunar swellings adjacent to and cridae of the Pacific Northwest of North America.
on either side of cardiac region.
Chelipeds equal. Manus of cheliped longer than Types. - The holotype, USNM 507767, and three
high, length to height ratio about 0.44, becoming paratypes, USNM 507768-507770, are deposited
with in the United States Museum of Natural higher distally; proximal margin convex, small, History,
and blunt Smithsonian D. C. blunt, spine at upper corner long, spine Institution, Washington,
lower at lower corner; margin developed into sinu-
collected Occurrence. - All of the were ous, serrate, keel, convex near proximal margin specimens
with in the late Eocene Hoko River and concave near distal margin; uppermargin from RB32, a locality
sharp keel ornamentedwith about five blunt nodes; Formation, located in the SW1/4, NW1/4, Sec. 4,
distal margin sinuous, oriented at about 120 de- T33N, R15W, Cape Flattery Quadrangle, 7.5' se-
with blunt Clallam several hundred gree angle to upper margin, broad, ries, Co., Washington,
the south shore ofthe Strait of swelling at midheight; outer surface of manus or- meters inland from
site. namented with tubercles on upper half, with weakly Juan de Fuca near a sanitary disposal
developed central keel, best developed distally; very differs from the and Remarks. - The new weakly developed keel between upper margin species closely
central keel, best developed centrally. related Anisospinos wahkiakumensis in several
has Fixed finger highest proximally and narrowed regards. Anisospinos berglundi a broader fronto-
distally, height to length ratio about 0.56; with orbital width to maximum width ratio (55 percent)
with than does A. wahkiakumensis The weakly hooked tipped; upper margin variably (40 percent). frontal sized, blunt denticles; outer surface with shallow ridges extending from the lateral spines are
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Table I. Measurements taken the dorsal of = = (in mm) on carapace Anisospinos berglundi. LI maximum length, L2 length to
maximum width = maximum = = = on carapace; Wl width, W2 frontal width, W3 fronto-orbital width, W4 posterior width.
Specimen Number (USNM) LI L2 W1 W2 W3 W4
507767 16.2 10.6 22.1 5.7 12.2 7.2
507768 14.7 10.3 20.7 5.2 11.4 5.9
507769 17 10.8 21.4 6.4 12 7.2
Table 2. Measurements (in mm) taken on the manus of the Glebocarcinus Nations, 1975
chelipeds of the holotype of Anisospinos berglundi, USNM
507767.111= maximum of LI = maximum height manus, length Cancer Nations, 1975, (Glebocarcinus) p. 22; Karasawa, 1990, ofmanus including fixed finger, L2 = length of fixed finger, L3 7, 1, 4a-c, 7a-b, 10-14; Davie, 1991, 494; - p. pi. figs. p. Sakumoto, length ofmovable finger, H2 height of fixed finger, H3 = and Karasawa, Takayasu, 1992, p, 447, pi. 60, figs. 5a-c, pi. height of movable finger. 61, fig. 1; Karasawa, 1993,p. 49, pi. 9, figs. 2-3, pi. 10, fig. 3;
7. Karasawa, 1997, p, 45, pi, 11, figs. 2, HI LI L2 L3 H2 H3
Left 6.5 14.7 4.7 N/A - 2.7 3.0 Type Species. Cancer oregonensis Rathbun, 1898 Right 6.4 15.0 5.4 7.7 2.8 3.0 by original designation.
much better developed in.A. wahkiakumensis. Aniso- Diagnosis. - Carapace length about three-quarters
spinos berglundi has more evenly spaced frontal maximum width. Front not produced beyond orbits;
spines ofabout the same size, while in A. wahkiaku- with five spines including inner-orbital spines, inner
mensis, the middle three spines are more closely three spines closely spaced, medial spine at lower
spaced. The anterolateral spines in A. berglundi level than other frontal spines; frontal region axially
are longer and better differentiated from each other sulcate, sulcus extending from position of medial than those ofA. wahkiakumensis, and additionally, frontal spine. Fronto-orbital width about 46 percent
the and sizes the between maximum orbits shapes of spines differ carapace width; moderate sized, the two species. In A. berglundi, the last antero- directed weakly anterolaterally, with two fissures.
lateral spine is long, attenuated, and directed weakly Anterolateral margin with nine spines, separated
to anterolaterally while in A. wahkiakumensis, that bases, directed forward, sharp, with granules or
spine is short and directed weakly posteriorly. The spinelets; spines two and three and four and five
seventh anterolateral ofA. is be Posterolateral spine berglundi nar- may paired. margin nearly straight
row and short, while in A. wahkiakumensis, it is or weakly concave, rimmed, with two small spines. broad and triangular in shape. Anisospinos wahkia- Posterior margin rimmed, straight. Regions inflated, kumensis has dense granules, while A. berglundi densely granular, defined by broad grooves. is ornamented with and Manus of five sparse granules scattered chelipeds short; rows of tubercles
tubercles the and branchial on lower larger on gastric re- outer surface; margin smooth; upper gions. The two species differ in several aspects of margin with four granular, bluntprojections. Fixed the first perieopods. Anisospinos wahkiakumensis finger short; narrow at tip, with two rows of tuber- is rather ornamented on the chelae; the manus has cles; occlusal surface with large, blunt denticles.
much a larger keel and larger serrations on the lower Movable finger with hooked tip, markedly narrowed
ofthe the surface of the rows of one margin manus; outer manus distally; two granules, on upper mar- has better developed keels and a much larger spine gin and one on outer surface; occlusal surface with
the blunt denticles. at upper distal corner of the manus. Anisospinos very short, herglundi differs from A. odosensis in possessing broader, longer anterolateral spines, a broader or- Remarks. - Nations (1975) referred five species bital rim, and a much to the Glebocarcinus. Of Cancer more granular carapace. subgenus these,
(Glebocarcinus) tumifrons has been removed to
Anatolikos. Nations (1975) referred Cancer balssi
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the that to subgenus Glebocarcinus; species has on outer surface, keels granular or smooth; lower been with with smooth synonymized Platepistoma anaglyptum margin or granular keel; upper mar-
(Balss, 1922) as discussed by Davie (1991). Glebo- gin with numerous sharp spines, granular, or with carcinus allisoni Nations, 1975 is known only from smooth keel; distal margin extending at approxi- claw recovered from middle Miocene fragments mately 120 degree angle to upper margin. Fixed
with central keel and another to late Pliocene deposits from Oregon to California, finger on lower mar- and it is difficult to confirm a generic assignment gin; occlusal surface with sharp denticles. Mov-
Karasawa for this incomplete material. (1997) re- able finger with sharp spines, granules, or smooth
G. from Pliocene of keel occlusal ported amphioetus deposits on upper surface; surface with large,
Japan; however, this occurrence is based upon one sharp denticles. movable finger and the generic placement cannot
be confirmed. the fossil record of this - Therefore, Remarks. Species of the genus Metacarcinus share
is and two Recent of smooth dorsal genus questionable, only species possession a relatively carapace;
the nine are referrable to genus. or ten fissured or small, serrate or granular
The two species of the genus form a small, tightly anterolateral spines, and rimmed posterolateral
defined restricted the North be ornamented with group to Pacific rim. margins which may one spine.
Glebocarcinus from Alaska Of oregonensis occurs to the taxa assigned to the genusby Nations (1975),
M. been removed California, and G. amphioetus has been reported plebejus has as discussed for Can-
from Metacarcinus Roma- China, Korea, Japan, California, and Baja cer s. s. granti was referred to
California. The genus probably evolved in the North leon by Nations (1975); however, it is hereinplaced
Pacific Ocean and within Metacarcinus because it subsequently dispersed through- possesses fissured
out that region, never becoming a speciose group. anterolateral spines with sharp, serrate terminations,
a feature for the diagnostic genus.
The chelae of to this species assigned genus are
Metacarcimis A. Milne Edwards, 1862 of two major types. In Metacarcinus anthonyi, M.
danai, M. davidi, M. edwardsii, M. gracilis, M.
M. Cancer (Metacarcinus) Nations, 1975, p. 23; Williams, 1984, jenniferae, novaezealandiae, and M. granti, the
Karasawa and p. 351; Sakamoto, Takayasu, 1992, p. 447, pi. claws resemble those of Cancer because s. s. spp. 60, figs. 5a-c, pi. 61, fig. 1; Karasawa, 1993, p. 50, pi. 10, fig. they possess short mani and fingers typical of and 4; Berglund Goedert, 1996, p. 830, figs. 2, 3; Karasawa, Cancer s. s. and spined or smooth keels. The claws 2-3. 1997, p, 46, pi. 10, figs. of M. magister resemble those of Romaleon be-
cause they have mani and ornamented - long fingers Type Species. Cancer magister Dana, 1852 by with prominent spines. Therefore, the original designation. morphology
of the chelae is not for this How- diagnostic genus.
all of the are to - evef, species readily assignable Diagnosis. Carapace ovate, about two-thirds wider
this based upon dorsal carapace morphology, than long. Front with five spines including inner- g,enus
innerthree particularly the shape of the anterolateral spines. orbital spine, spines closely spaced; front The oldest verifiable occurrence ofMetacarcinus usually not produced beyond orbits. Fronto-orbital based dorsal is M. width about 0.26-0.34 maximum upon carapace morphology goe- carapace width;
orbits derti new species from Oligocene rocks of Alaska shallow, directed forward. Anterolateral mar- discussion of with (but see Metacarcinus sp. below). gin nine or ten spines; anterolateral spines Metacarcinus starri was described from early variable in form; small, sharp, and separated to Miocene rocks of the Pysht and Clallam forma- bases or small, sharp and fissured; spine margins tions of Washington, USA. Metacarcinus danai has simple, serrate, or granular. Posterolateral margins been reported from the Miocene of California, and rimmed, sometimes with one spine; carapace re- M. gions jenniferae was reported from the middle Pliocene poorly developed, smooth or ornamented with of California. Metacarcinus davidi is fine granules. represented
by carapace material and claw material from middle Manus of chelipeds long; usually with four keels
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Miocene to middle Pliocene rocks of California. Fronto-orbital width about 32 percent maximum
the known orbits directed The two Japanese species of genus are carapace width; circular, forward;
from dorsal material and include the two orbital inner fissure about mid-width carapace fissures, at
Miocene M. izumoensis and the Pliocene M. minuto- oforbit, outer fissure just proximal to outer orbital
serratus. Metacarcinus magister, M. gracilis, and spine. Anterolateral with nine M. anthonyi are known almost exclusively from margin convex; spines
claw fragments from Pliocene and Pleistocene rocks including outer-orbital spine, spines separated to with anterior and of the west coast of North America. Because the bases, concave margins convex
from each other oldest occurrence of the genus is in North America, posterior margins, separated by
it is that evolved in the North sharp, v-shaped notches, spines appearing to be suggested the genus
ornamented with First the outer- Pacific, as early as the Oligocene, and subsequently granules. spine orbital spine, broad, directed forward, curving dispersed throughout the North Pacific Ocean by
similar distribution weakly toward axis; second spine curving axially, the Miocene. The genus has a
that ofCancer s. in modern directed forward, smallest of anterolateral spines; as s. oceans, inhabiting third broad, directed forward, axi- the northeastern, southeastern, and southwestern spine curving fourth directed fifth Pacific and the North Atlantic Oceans. The similar ally; spine broad, forward; spine
about as broad as fourth, directed anterolaterally; geographic and chronologic distributions of Meta- sixth spine about as large as fifth, directed antero- carcinus and Cancer s. s. suggest that they are laterally; seventh spine broadest of anterolateral especially closely related. spines, directed anterolaterally; eighth spine about
as broad as fourth-sixth, directed anterolaterally;
ninth spine small, triangular, sharp, directed later- Metacarcinus goederti new species ally. (Figs. 4-5) Posterolateral margin sinuous, with granular rim,
initially nearly straight, then concave posteriorly. Diagnosis. - Carapace wider than long, widest at Posterior margin nearly straight, rimmed, 38 per- position of last anterolateral spine located about cent maximum carapace width. 60 distance percent posteriorly on carapace; cara- Protogastric region triangular in shape, weakly surface smooth pace or finely granular, regions developed, flattened near frontal region, inner and weakly defined; orbits large for genus; anterolateral outer margins convex. Mesogastric region narrow margin with nine sharp spines separated to bases, anteriorly, broadened posteriorly; margins of an- ornamented with fine granules; posterior margin terior process concave; posterior margin weakly entire, rimmed. convex. Urogastric region not well-differentiated
from other gastric regions, with concave lateral
- wider than Description. Carapace long, (ength anterior margins, margin concave, posterior margin about 68 percent maximum width, widest at posi- nearly straight. Cardiac region rounded triangular, tion of last anterolateral located about 60 spine directed rounded apex distally, two swellings an- distance percent posteriorly on carapace; carapace teriorly, smaller rounded swelling posteriorly. In- regions weakly inflated, delimited by shallow testinal region flattened, not well-developed. surface to be smooth grooves; appearing relatively branchial Hepatic and regions very weakly in- or finely granular; vaulted both longitu- carapace flated, poorly developed. Metabranchial region with dinally and transversely. semi-lunar swellings adjacent to and on either side Front about 16 maximum width, percent carapace of cardiac region. not projected beyond orbits. Front with five spines Pterygostomial region finely granular, with two including inner-orbital spines, medial spine long- subhepatic grooves. Suborbital margin appearing est, narrow, lateral spines short, blunt, inner- narrow; to have broad, triangular notch. orbital spines separated from lateral spines by broad, Remainder of venterand appendages not known. u-shaped emargination, shortest of frontal spines, broadly triangular. Measurements. - Measurements (in mm) taken on V / \ ?
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4. Dorsal of USNM 507772. Dorsal of USNM Fig. Melacarcinus goederti n. sp. 1, carapace paratype, 2, carapace holotype, 507771.
specimens of Metacarcinus goederti are given in Types. - The holotype, USNM 507771, and four
Table 3. Position and orientation of measurements paratypes, USNM 507772-507774, are deposited
States National taken are illustrated in Fig. 5. in the United Museum of Natural
Flistory, Smithsonian Institution, Washington, D.C.
- James L. Etymology. The trivial name honors
Goedert, Gig Harbor, Washington, who has made Occurrence. - The specimens were recovered from a significant contribution to the study of the the BearLake Formation, questionably of Oligocene
Cancridae of North USA. The of the Pacific Northwest age, Alaska, holotype, USNM 507771,
27 America. was collected from Sec. or 34, T48S, R69W,
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Metacarcinus goederti differs from all other
referred to the in species genus Metacarcinus pos-
sessing very well-differentiated anterolateral spines.
Metacarcinus plebejus, M. novaezealandiae, M.
and all anterolateral davidi, M. anthonyi possess
spines that are fissured and not separated to their
bases as in M. goederti. Metacarcinus anthonyi
also has much smaller orbits than does M. goederti.
Metacarcinus magister has a more granular cara-
shallower and smaller anterolateral pace, orbits,
spines ornamented with spinelets or large granules
than does M. in which the anterolateral goederti ,
spines are simple or ornamented with granules.
Metacarcinus danai has smaller and less well-dif-
ferentiated spines and smaller and shallower orbits
than does M. Metacarcinus minutoserratus Fig. 5. Line drawing of Metacarcinus goederti showing the goederti. position and orientation of measurements taken. has complex and serrate anterolateral spines, while
those of M. goederti are simple and granular. Me-
tacarcinus starri has small anterolateral spines
Table 3. Measurements (in mm) taken on specimens of ornamented with tiny serrations, whereas the an- Metacarcinus goederti. LI = maximum length, L2 = length to terolateral spines of M. goederti are larger and maximum width on carapace; W1 = maximum width, W2 =
to M. the frontal width, W3 = fronto-orbital width, W4 = posterior width. granular. Additionally, contrary goederti,
medial frontal spine of M. starri is the smallest of Specimen the frontal spines, and the lateral medial teeth are Number largest. Finally, in M. starri, the dorsal carapace (USNM) LI L2 W1 W2 W3 W4 while in is smooth, M. goederti, the carapaceregions
507772 39.7 23.3 52.7 8.2 16.6 21.4 are weakly defined. Metacarcinus izumoensis has 507771 35 20.3 53.2 8.8 16 N/A smaller, less produced anterolateral spines than does 507773 31.7 18.6 43.2 6.2 15.2 15.7 M. and exhibits goederti more pronouned carapace
ornamentation.
Port Mollcr (D-1) Quadrangle, USGS M8170. The remainder of the material was collected from the
?Metacarcinus sp. Milky River section of the Bear Lake Formation, (Fig. 6) Sec. 27 and 34, T48S, R69W, Port Moller (b-l)
Quadrangle, USGS M8171. USGS M8171 is slightly - Material Examined. One claw, referredto GHUNLPam 16803, higher stratigraphically than USGS M8170. is deposited in the Paleontology collection of the Facultad de
Ciencas Exactas Universidad Nacional de La y Naturales, Pampa,
- Metacarcinus Remarks. The new species goederti Santa Rosa, La Pampa, Argentina.
based is placed within Metacarcinus upon posses-
sion Occurrence. - The ofnine anterolateral spines including the outer- specimen was collected from
25 orbital spine; anterolateral spines that are distinctly the Centinela Formation, on the Estancia de separated and ornamented with granules; five frontal Mayo, about one km east of the locality described spines with the middle three closely spaced; a front below for Notocarcinus sulcatus new species. The not extending beyond the orbits; rimmed postero- claw was collected just below the volcanic tuff
and N. is lateral posterior margins; and indistinct cara- described for sulcatus and therefore middle
the lacks in pace regions. Additionally, new species Eocene age. posterolateral spines, which is typical of many species of Metacarcinus. Remarks. - One claw fragment was recovered from
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Diagnosis. - Small cancrid with projecting front,
anterolaterally directed orbits, and undulose cara-
axial pace surface with depressed and hepatic ar-
eas separated by broad protogastric ridges.
Description. - Carapace diamond-shaped, length
maximum width. about 73 percent carapace Front
projecting well beyond orbits, with five projec-
tions including innerorbital spine, innerthree pro-
jections truncated, closely spaced, separated by
shallow fissures; frontal region deeply and broadly
view of 6. Metacarcinus outer cheliped, GHUNLPam sulcus between inner or- Fig. sp. , sulcate, spanning region
16803. bital spines. Fronto-orbital width about 44 percent
maximum carapace width; orbits shallow, broad,
middle Eocene rocks of the Centinela Formation with two fissures, directed anterolaterally. Ante-
near the locality from which the specimens of rolateral margin gently and smoothly curved; eight sulcatus the Notocarcinus were collected. Because anterolateral spines, including outer orbital spine,
claw is than the dorsal as large or larger carapace spines separated by fissures, truncated at tips,
of specimens of N. sulcatus, it is not referred to margins of spines granular. Posterolateral margin
that Taxa referred to the Cancridae do not rimmed species. very weakly concave, at least in the pos-
exhibit such in size relative to the a disparity claw terior half, not spined. Regions weakly developed,
size of the most in carapace. prominently the posterior axial area; he-
The claw material is referred to the questionably patic, frontal, and epigastric areas depressed.
genus Metacarcinus. The manus is short and high;
ornamented four keels the - The with or five granular on Etymology. name was derived from the Greek
outer surface and a granular keel on the lower “notos,” south, and “carcino,” crab, in reference
and it have a keel with a few the of the in the southern margin; appears to to occurrence genus
the All of these large prominences on uppermargin. hemisphere.
are characteristics typical of the mani of species
of both Cancer s. s. and Metacarcinus. The material - Occurrence. The genus is described from speci-
is referred Metacarcinus the distal to because margin mens collected at a single locality on the Estancia of the makes about 120 with manus a degree angle 25 de Mayo, near Calafate, Argentina. A detailed
the most of Cancer upper margin. In species s. s., description of the locality is given below.
that angle is about 90 degrees. If this material is in
fact of the then Remarks. - representative genus Metacarcinus, Placement of this new genus within
the of the is extended into the geologic range genus Cancrinae can be made with confidence. The
the middle Eocene. this is wider than the frontal bears Furthermore, might suggest carapace long, region
a southern hemisphere origin for the genus with five projections, the orbits exhibit two well-devel- the North Pacific Ocean subsequent dispersal into oped fissures, the anterolateral margin is divided
the into blunt by Oligocene. Confident generic assignment eight projections separated from one
awaits the of dorsal material. another distinct and the rimmed discovery carapace by fissures, pos-
terolateral margin lacks spines. However, several
characteristics serve to distinguish the genus from
others within the The frontal Notocarcinus new genus subfamily. margin
projects well in advance of the orbits, and the three
- sulcatus new central of the front blunt and Type species. Notocarcinus spe- projections are sepa-
cies by original designation. rated by fissures whereas the outer projections, the
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inner orbital spines, are more distinctly separated vaulted laterally and transversely with depressed and pointed. The orbits are directed anterolaterally frontal and hepatic areas, other areas defined as
inflated and the frontal and epigastric regions, are deeply regions separated by broad, indistinct depressed and bordered by protogastric ridges depressions. extending from the orbits posteriorly and axially. Front generally straight, projected slightly in
Lateral to these ridges, the hepatic regions are advance of orbital regions, with two short, narrow
that the entire front half of the fissures three blunt front defined depressed so carapace defining teeth; has an undulose surface. Whether this character is laterally by distinctly pointed corner extending
in fact diagnostic of the genus or is a species-level posterolaterally to fissure at inner orbital tooth,
await additional frontal width about maximum trait must discovery of species 19 percent carapace within the the of the width. Orbits directed with genus. Regardless, placement anterolaterally two specimens within a unique genus seems certain. closed fissures in outer halfof orbit and bounding
The occurrence of Notocarcinus in middle Eocene rounded projection, fronto-orbital width about 45
rocks in is Glaessner maximum width. Anterolateral Argentina significant. (1969) percent carapace
the of the Cancridae extend from shorter than reported range to margin posterolateral margin, distinct,
middle Eocene to Recent; however, the Cancrinae weakly upturned, gently convex in dorsal aspect, were known only from Miocene to Recent occur- with eight blunt, undivided lobes, including outer
The of the Cancrinae orbital lobes from another rences at that time. range spine; separated one by
Eocene has been extended into the middle as re- closed fissures. Posterolateral margin straight to corded by Notocarcinus, which is the oldest known slightly concave; posterior margin straight, wider occurrence of the subfamily. Perhaps even more than front, posterior width about 25 percent maxi-
the Cancrinae is width. Prominent rim significantly, currently widely mum carapace marginal distributed within the Pacific Ocean basin and in borders posterior half of posterolateral margin and
from the North Atlantic. No cancrids are known posterior margin. modern seas along the east coast ofSouth America. Frontal region broadly sulcate, bounded by
In the fossil record, Beurlen (1958) reported protogastric ridges converging posteriorly from
tuberculatus from the inner orbital to elevated ?Cyclocancer upper Oligo- spine mesogastric region
lower cene or Miocene of Brazil and van Straelen that is slightly longer thanwide; metagastric region
(1937) described Parapirimela angolensis from weakly distinguished from mesogastric by subtle
West Africa. The Brazilian material was originally transverse depression. Cardiac region circular, assigned to the Atelecyclidae. It may, in fact, be a bearing two small nodes at about midlength. In- cancrid; however, the assignment remains in doubt. testinal area indistinct. Carapace surface granular,
The somewhat African occurrence is probably not a cancrid most distinctly on elevated regions and as discussed above. Thus, the occurrence of Noto- smoother in depressions. carcinus sulcatus from Argentina is the first wtell- Epistome weakly convex laterally, surface documented cancrid, fossil or extant, known from smooth. the South Atlantic region. Remainder of venter, abdomen, and legs not
known.
Measurements. - Notocarcinus sulcatus new species All measurements were taken on
(Figs. 7-8) the holotype, GHUNLPaml6801. The paratype is
incomplete. Maximum length, 22.9 mm; length to
maximum width - on 12.2 maximum Diagnosis. As for genus. carapace, mm; 31.4 frontal 6.1 fronto- width, mm; width, mm;
orbital 14.2 7.8 Description. - Small cancrid; elongate ovoid out- width, mm; posterior width, mm. line, wider than long, length about 73 percent
maximum widest about - The trivial name alludes the de- width, carapace 53 per- Etymology. to
distance exhibited on the cent posteriorly on carapace; strongly pressed hepatic regions specimens
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7, Notocarcinus sulcatus and Dorsal of GHUNLPam 16801. Dorsal of Fig. n. gen. sp. 1, carapace holotype, 2, carapace paratype,
GHUNLPam 16802.
the Estancia 25 de Mayo, Latitude 50°30.716’S,
km Longitude 72°!5.303’W, ca. 12 south from
Calafate, Santa Cruz Province, Argentina. The
deposits are dated at 46 ma on the basis ofa promi-
nent volcanic tuff, exposed just above the site from
which the crabs were collected (Casadio et al.,
2000).
Remarks. - The two specimens are moderately well
of preserved in terms the dorsal carapace; cuticle
preserved on the anterior two-thirds of the cara-
confirms that the ornamentation is limited pace to
fine The has a characteris- very granules. species
tic diamond-shaped outline, because the antero- Fig. 8. Line drawing of Notocarcinus sulcatus. lateral margins are somewhat straighter than in most
meiribers of the family. However, the undulose
nature of the anterior of the and that serve to distinguish this species from all other portion carapace
cancrids. the relatively distinct definition of the posterior
regions are certainly the most diagnostic features
- of It Types. The holotype, GHyNLPam 16801, and this crab. cannot be mistaken for any other cancrid. paratype, 16802, are deposited in the
Paleontology collection of the Facultad de Ciencias
Exactas Universidad Nacional de La y Naturales,
Pampa, Santa Rosa, La Pampa, Argentina. Platepistoma Rathbun, 1906
- Rathbun, 876. Occurrence. The two specimens were collected Plalepistoma 1906, p.
Cancer 4B. (Platepistoma) Takeda, 1977,p. 35-38, fig. from the Centinela Formation, middle Eocene, on
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- Type Species. Platepistoma macrophthalmous Islands, French Polynesia, New Caledonia, Tasman
Rathbun, 1906 by original designation. Sea, the Seychelles, and the Sala-y-Gomez Ridge
of the southeastern Pacific (Davie, 1991). This
Diagnosis. - Carapace hexagonal, three-fourths as suggests that the genus may have evolved during
Front long as wide. projected slightly beyond orbits; the Miocene in Japan and subsequently dispersed
front with five poorly developed, short, spines throughout the Indo-Pacific region.
including inner-orbital spines, medial spine at lower
level than other frontal spines. Fronto-orbital width
from 38 to 44 maximum percent carapace width; Romaleon Gistl, 1848
orbits shallow, very small, directed forward, with
two fissures. Cancer Anterolateral margin with eight or (Romaleon) Nations, 1975, p. 23; Davie, 1991, p. 508; Berglund and Goedert, 1992, 4, figs. 2-16; nine spines; spines separated tobases, simple, small, p. Karasawa, 1993,
p. 50, pi. 10, fig. 6; Karasawa, 1997,p. 45, pi. 10, fig. 6, 8, reduced. Posterolateral margin nearly straight, entire
or with one spine, rimmed. Posterior margin straight,
Type Species. - Cancer gibbosulus Rathbun, 1898. rimmed. Regions well defined, ornamented with
densely spaced tubercles; regions defined by deeply
- 60 Diagnosis. Carapace to 75 percent as long as excavated, smooth grooves. wide. Front slightly produced beyond orbits; five Manus of chelipeds short; outer surface with four frontal spines including inner-orbital spines, inner or five rows of granules; lower margin smooth; three very medial at with several short spines closely spaced, spine upper margin spines or granular. lower level than other frontal spines; frontal region Fixed finger with two rows of granules, occlusal axially silicate, sulcus extending from position of surface with large, blunt denticles. Movable finger medial spine. Fronto-orbital width 29 to 44 percent smooth, occlusal surface with very short, blunt den- maximum carapace width; orbits moderate sized, ticles or nearly smooth. shallow, directed forward. Anterolateral margin with
nine separated to - spines, spines bases, sharp, curving Remarks. Davie (1991) provided a thorough forward. Posterolateral margins concave, rimmed, discussion of this genus, a small, tightly defined sometimes with one Posterior and offered several which dif- spine. margin nearly group, means by to straight, rimmed. Regions typically moderately ferentiate members of Platepistoma from Cancer defined, sometimes with granules or transverse s. l., which have been expanded upon in the diag- ridges. nosis. The Japanese species Cancer? imamuraehas Manus of outer surface with inflated ornamented with chelipeds elongate; carapace regions densely four-six granular keels; lower smooth or the margin spaced granules; deep grooves separating cara- with with granular keel; upper margin numerous pace regions; short, poorly developed frontal spines; sharp, distally directed spines. Fixed finger with and small, poorly developed anterolateral spines. two granular keels, occlusal surface with large, blunt Cancer? imamurae differs from other members of denticles. Movable surface with in finger long; upper Platepistoma possessing several small spines on directed distally spines or granules; occlusal surface the posterolateral margin; however, this is the only with large, blunt denticles. significant difference between Cancer? imamurae and the other members of the genus. Therefore, it
Remarks. - Members of the Romaleon genus are seems most prudent to place Cancer? imamurae united by their possession of singular, anteriorly- within Platepistoma instead of erecting a mono- curving anterolateral spines that are separated to for this specific genus species. their bases. Most members of the have elon- genus The two fossil of this species genus, Platepistoma gate mani and fingers of the cheliped, and those imamuraeand P. kaedei, are known from Miocene elements are typically ornamented with or of spines deposits Japan. Modern members of the genus sharp granules. inhabit the Indo-Pacific region including occur-
Cancer nadaensis Sakai, 1969 was referred to rences in Hawaii, Japan, Madagascar, the Reunion Cancer (Cancer) by Nations (1975). That species
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was placed within the subgenus Romaleon by two fissures; anterolateral margin typically with
Karasawa (1990); it is herein placed within Roma- several anterolateral spines; spines well-developed, leon. Kato three dorsal often bifurcate (1996) figured carapace separated to base, or trifurcate; specimens (figs. 5-5, 5-6b, 7) as members of the posterolateral margin typically coarsely granular new species Cancer sakamotoi, here referred to or with well developed spines; posterior margin the Romaleon. Examination of these genus figures nearly straight, granular or spined; carapace regions indicates that the holotype specimen for R. sakamo- typically moderately well-developed, ornamented toi, illustrated in fig. 7 (Kato, 1996), is indeed with dense granules or with discrete, large spines. distinct. the illustrated However, carapace in fig-
5-5 is based - ure referrable to Romaleon sanbonsugii Remarks. The Lobocarcininae is a small sub-
the of the anterolateral and the of three of which upon shape spines family comprised genera, two marked curvature of the anterolateral margin. The are monospecific. Miocyclus differs from the other
in differs from the specimen figured 5-5 holotype two included genera in possessing fissured antero- of C. in tri- lateral sakamotoi possessing small, broadly spines and a smooth, entire posterolateral
that are directed it number of angular, spines, anterolaterally, margin. However, possesses an even while the holotype specimen has narrow,attenuated frontal spines, a diagnostic character of the sub- spines that are separate to their bases and curve family. Therefore, it is placed within the Lobocar- weakly anteriorly. cininae, although this placement can be considered
The oldest known occurrences of Romaleon, to be provisional because the specimens have not based upon dorsal carapace morphology, are R. been examined.
dereki from the middle Miocene California and All of ofthe genera within the Lobocarcininae have
R. R. middle their earliest in known sakamotoi and sanbonsugii from occurrence or are solely
Miocene rocks of Japan. Only one other fossil from the Tethys, and in fact, only one species of
occurrence based upon dorsal carapace material is Lobocarcinus, L. pustulosus Feldmann and Fordyce, known from North America, R. urbanus, from 1996, is extralimital. This suggests that the sub-
in the the Pliocene rocks of California. Fossil occurrences family arose Tethyan region, as early as based claw include R. R. middle and into upon fragments jordani, Eocene, subsequently dispersed branneri, R. antennarius, and R. polyodon from New Zealand waters.
Pliocene to Pleistocene deposits of the west coast of America. The have North genus appears to
the evolved in the North Pacific Ocean as early as Lobocarcinus Reuss, 1857
Miocene and subsequently dispersed throughout
the - North Pacific Ocean. One modern species ranges Type species. Lobocarcinus paulinowuerttem-
into Pacific the southeastern region. The genus has burgensis von Meyer, 1847. a similar geographic distribution to that of Cancer
and - Metacarcinus, suggesting a close relationship Diagnosis. Carapace much wider than long. Front between these three genera. projected weakly beyond orbits; four to six frontal
spines including inner orbital spines. Fronto-orbital
Lobocarcininae 1930 width from 30 40 maximum Subfamily Beurlen, to percent carapace
width; orbits round, with two fissures, directed
Included genera. - Lobocarcinus Reuss, 1857; forward; orbits rimmed. Anterolateral margin usu-
in Miocyclus Muller, 1979; and Tasadia Muller ally with seven lobes; lobes typically sharply bi- Janssen and Muller, 1984. furcate or trifurcate. Posterolateral margin nearly
straight, with several spines of varying size. Pos-
Diagnosis. - Carapace wider than long; front usually terior margin nearly straight, sometimes with spines.
with even number of from four to spines, ranging Regions moderately well-developed, often orna- six and ifodd number of spines, central spine small mented with several large nodes. Chelipeds slender and at lower level than other spines; orbits with and equal.
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- The is from the Remarks. Six species are referred to the genus genus known only fossil record,
Lobocarcinus. Ail are known from the Tethyan occuring in middle to late Eocene rocks of Egypt, region with the exception of L. pustulosus, known Hungary, and India; Miocene rocks ofNew Zealand; from the Miocene of New Zealand (Feldmann and and Miocene to Pliocene rocks ofthe Mediterranean
A discussion rim and Northern Feldmann and Fordyce, 1996). complete of the group Europe. Fordyce
and that the evolved in the was provided by Anderson Feldmann (1995), (1996) suggested genus who revised the and described and from that genus a new spe- Tethys subsequently dispersed region cies. Lobocarcinus pentanodosus Collins and Bar- into New Zealand. Members of this genus are the
Eocene In- earliest of the ber, 1998, was reported from rocks of known members subfamily.
donesia; however, that species has fewer antero- lateral spines than do species ofLobocarcinus and the spines of L. pentanodosus are singular, not Miocyclus Müller, 1979 bifurcate or trifurcate as in authentic Lobocarcinus.
The posterolateral margins ofL. pentanodosus are Type species Miocyclus bulgaricus Müller, 1979. entire, while those of authentic Lobocarcinus are
The ornamentation ofL. - wider than spined. carapace pentano- Diagnosis. Carapace long, flattened, dosus differs from that of authentic Lobocarcinus. ornamentedwith small, dense granules; orbits with
Typical members of Lobocarcinus possess large two fissures; front weakly projecting beyond orbits,
the and rather well defined granules on carapace with six frontal spines including inner-orbital spines;
Lobocarcinus has first carapace regions. pentanodosus anterolateral margin crispate, with nine spines; poorly defined regions and transverse ridges on eight fissured, with blunt margins; last spine sharp, the protogastric regions. The transverse ridges and produced; posterolateral margin entire, nearly the possession of five, singular anterolateral spines straight; posterior margin weakly convex.
be suggest that L. pentanodosus may a portunid.
sismondae is here referred to the - Cancer genus Remarks. Miocyclus is a monospecific genus
Lobocarcinus based upon its possession of distinctly known from the Miocene ofBulgaria. It was origi- lobed anterolateral margins, spined posterolateral nally placed within the Atelecyclidae by Muller
the overall of the and margins, and shape carapace (1979) but was later moved to the Cancrinae based
and carapace regions (Bonfiglio Donadeo, 1982). upon possession of wide mcsogastric regions
The to be ornamented with The is here within carapace regions appear (Muller, 1984). genus placed
as is of of Lobocarcinus. the Lobocarcininae based of nodes, typical many species upon possession an
The difference between L. sismondai and other only even rather than odd numberof frontal spines. The
of Lobocarcinus is that it five species possesses genus is easily distingushed from other members
frontal spines, including an axial spine instead of of the Lobocarcininae because it lacks well-devel- an even number as is typical of the genus) and oped bifurcate or trifurcate spines on the antero- subfamily. This constitutes a major difference; lateral margin and has a smooth, entire posterolat-
all characters however, other carapace clearly place eral margin.
L. sismondai within Lobocarcinus. The axial spine is smaller than the other spines and placed at a
members of lower level, a condition seen in many Tasadia Müller in Janssen and Müller, 1984 the Cancrinae and which could represent conver-
It is also that Cancer Cancer gence. recognized illyricus Bittner, 1884,p. 27, pi. I, figs. 8-9; Lorenthey,1897, p,
Bittner, 1893 and C. deshayesii A. Milne Edwards, 159-160, 167-168.
Atelecyclus Lorenthey. Lorenthey in Lorenthey and Beurlen, 1861 L. are synonymous with sismondai (Glaessner, 1929,p.156-158, pi. 9, fig. 9; gripp, 1969, pp. 86-87, pi. 3, fig, 1929; Muller, 1984; Collins and Fraaye, 1991). I.
of these taxa were not seen but Specimens by us, 15 Cyclocancer (Lorenthey). Beurlen, 1958, p. the of these taxa is well-demonstrated Cancer 6. synonymy (Glebocarcinus) Karasawa, 1990, p.
in the cited works.
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Type species. — Cancer carniolica Bittner, 1884. and the frontal spines are much better developed
than in Tasadia. For all of these reasons, Tasadia
Diagnosis. - Carapace wider than long, L/W about is retained as a distinct genus. As far as is known,
0.60. Front not projected beyond orbits, four or Tasadia is a monospecific genus restricted to the five frontal spines. Fronto-orbital width to width Miocene of Central Europe. Cancer bittneri Toula, ratio about 30 percent; orbits small, shallow, di- 1904 was recognized to be synonymous with T. rected forward, with two fissures. Anterolateral carniolica by Muller (1984). margin convex, ornamented with eight or nine tri- angular spines, with granules or spinelets, each separated to bases. Posterolateral margin nearly Discussion straight, granular, with several small spines. Pos-
The Cancer l., and terior margin rimmed, nearly straight. Regions well biogeographic distribution of s. /.,
defined, ornamented with densely spaced tubercles; by implication, the Cancridae, has been addressed
defined regions by smooth, deep grooves. on numerous occasions (Ekman, 1953; Nations,
Manus of chelipeds with at least four granular 1975; 1979; Carvacho, 1989; Newman, 1991).
keels on outer surface; upper margin appearing to Ekman (1953), Nations (1975; 1979), and Carvacho
be lower ornamented with sharp spines; margin (1989) suggested that the genus had a primarily
smooth. Fixed with keels finger two granular on North Pacific Ocean origin and subsequently dis-
outer surface. Movable finger with sharp granules persed throughout the Pacific Ocean by following
on upper margin; at least one row of granules on the continental shelves; to New Zealand via Ant-
outer surface. arctica; and to the Atlantic Ocean by crossing the
Arctic polar ocean. Newman (1991) suggested that
Discussion. - that this relict dis- Davie (1991) suggested genus Cancer s. l. displayed a amphitropical
Plate- exhibited sufficient similarity to the genus tribution in the temperate North Pacific and South
that it be with whereas pistoma may synonymous it, Pacific Oceans. This conclusion was based on the
Karasawa Tasadiawith Cancer belief that l. the sole (1990) synonymized Cancer s. was extant genus
s. l. Examination of illustrations and casts of spe- of the family. The extinction of several tropical to
cimens of Tasadia in Janssen and Muller, in the Eocene Miocene (Muller subtropical genera to was
that it is indeed distinct 1984, pi. 5) indicates a cited as evidence that the genus, and by implication,
genus. Tasadia differs from members of Platepis- the family, arose and subsequently became extinct
toma in possessing distinct, large, triangular ante- in the tropics, resulting in a formerly Tethyan dis-
of Cancer rolateral spines, while the anterolateral spines tribution for s. l. and the family. However,
Platepistoma are small and reduced. Additionally, this hypothesis is no longer tenable for several
Newman Tasadia possesses a multi-spined posterolateral reasons. The genus Cancer sensu has been
margin, while that of Platepistoma typically only redefined; for example, Newman (1991) consid-
four the has one spine. The regions of Tasadia are not as ered species of all of former subgenera of
the Cancer members of the densely ornamented, and shape of the regions as well as genus Platepis-
differs in Tasadia from those of Platepistoma. toma in his discussion of the genus. The redefinition
is from Glebocar- results in different distribu- Tasasdia also distinguishable of these genera a very
the cinus, members of which typically have narrow, tional pattern for Cancer s. s. as well as four
anterolateral that directed ante- additional Newman considered spines a're weakly genera (1991) as
riorly and sometimes ornamented with spinelets. members of Cancer s. l. Furthermore, Newman
The anterolateral spines of Tasadia are short, broad, (1991) cited the appearance and extinction of Para-
and triangular and are ornamented with large gran- pirimela in the Miocene of Angola as evidence of
ules. The of Tasadia are not distribution of the Cancridae. carapace regions nearly a formerly tropical
well that as developed and are more sparsely yet more However, genus is probably not a member of
Glebocarcinus. coarsely ornamented than those of the family, and in any case, its familial placement
The in Glebocarcinus orbits are better developed cannotbe verified. The revision of each of the genera
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Cancridae makes it in with into the south- within the possible this report Ocean, one species ranging
to recast the biogeographic history of the family, eastern Pacific region. These distributions suggest
the Ocean two included subfamilies, and each genus (ge- that the North Pacific is the area of origin
neric discussions above). and subsequent dispersal for almost all of the
The earliest known confirmed of the cancrine occurrences genera.
family are Lobocarcinus of the Lobocarcininae from
the middle Eocene of Egypt and Notocarcinus of
the Cancrinae from the middle Eocene of South Acknowledgements
America. Anisospinos is known from late Eocene
rocks of Washington, and Metacarcinus makes its The specimens of Anisospinos berglundi were collected and
rocks ofAlaska. donated by R, E. Berglund, Bainbridge Island, Washington. first appearance in Oligocene (?)
Specimens of Metacarcinus goederti were collected by L. Several make their first in the genera appearance Marincovich and donatedby Charles L. Powell II, both of the Miocene, including Cancer s. s. from Japan, Ana- United States Geological Survey, Menlo Park, CA. Locality tolikos from Platepistoma from Japan, Ro- Japan, information for M. goederti was supplied by Powell. The from and from maleon Japan California, Miocyclus holotypes of Anisospinos wahkiakumensis and Metacarcinus
starri R. Thomas Bulgaria, and Tasadia from Romania and Belgium; were kindly loaned by Eng, Burke Memorial Washington State Museum, Seattle. The holotypes of Branchio- the latter two genera are members of the Lobocar- lambrusaltus and several cancrid taxa were graciously loaned cininae and the rest are members of the Cancrinae. by W. Blow, United States National Museum of Natural History, It is difficult to determine when and where the Washington, D.C. F. Collier, Museum ofComparative Zoology,
family first appeared because the earliest known I larvard University, Boston, MA, loaned the holotype ofCancer
occurrences from Eocene rocks are known from proavitus. H. Kato, Natural History Museum and Institute, Chiba,
Japan, and H. Karasawa, Mizunami Fossil Museum, Mizunami, each of the two subfamilies. This suggests that the Japan,provided helpful discussions about Japanesecancrids and family arose sometime before the middle Eocene. provided access to specimens of the Japanese taxa discussed further when However, speculation upon exactly here at their respective institutions.The manuscript was greatly and where this occurred is unwarranted until more improved by the thoughtfulreviews ofJoe S. H. Collins, London,
fossil material is recovered. England, and the late Austin B. Williams, United States National
Several from these Museum ofNatural History, Washington, D.C. Travel to examine patterns emerge occurrences. Japanese fossil material was funded by National Geographic the subfamilies have had dif- Clearly, two very Society Grant 6265-98 to Feldmannand Schweitzer. Travel and ferent evolutionary and dispersal histories. The field work to collect Argentinian materialwas funded by National
Lobocarcininae has a distribu- primarily Tethyan Geographic Society Grant 5588-95 to Feldmann.
discussed tion, as has already been above, since
all but of known from one species one genus are
the The Cancrinae have Tethys region. appear to References
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Weber F. 1795. Nomenclator entomologicus secundum 1962), as Cancer; P. kaedei (Karasawa, 1990)new comb.,
entomologiamsystematicum ill. Fabricii, adjectis speciebus as Cancer (Glebocarcinus);kiribatienseP. Davie, 1991;
recens detectis et varietatibus. Chilonii (Kiel) et Hanburgi, Platepistoma macrophthalmous Rathbun, 1906; P. nanum
1-171. Davie, 1991; P. balssi (Zarenkov, 1990), as Cancer; P.
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Smithsonian Institution Press, 1-550. Cancer; R. branneri (Rathbun, 1926), as Cancer; R. dereki
Yokoya Y. 1933. On the distribution of Decapod Crusta- Nations, 1975, as Cancer (Romaleon); R. gibbosulus (Rathbun, ceans inhabiting the Continental Shelf around Japan, based 1898), as Cancer; R. jordani (Rathbun, 1900), as
nadaensis upon the Materials collected by S. S. Soyo-Maru, during Cancer; R. (Sakai, 1969) new comb., as Cancer;
the 1923-1930. R. R. sakamotoi years J. Coll. Agriculture Tokyo Imperial polyodon (Poeppig, 1836), as Cancer; (Kato,
University 12: 1-225. 1996) new comb., as Cancer; R. sanbonsugii (Imaizumi,
Zarenkov NA. 1990. Decapods (Stenopodidea, Brachyura, 1962),new comb,, as Cancer; R. urbanus (Rathbun, 1917),
Anomura) of the Nacza and Sala-y-Gomez underwater as Cancer.
ridges. Trans. P. P. Shorshov Inst. Oceanol. Ac. Sci. USSR II. Family Cancridae,Subfamily Lobocarcininae
124: 218-244. Genus Lobocarcinus: Lobocarcinus aegypticus Ldrenthey, 1909;
L. indicus Glaessner, 1933; L. lumacopius Anderson and
Accepted: 5 June 2000 Feldmann, 1995; L. paulinowuerttemburgensis von Meyer,
1847;L.pustulosus Feldmann and Fordyce, 1996;L. sismon-
dai von Meyer, 1843, new comb., as Cancer.
Genus Miocyclus: Miocyclus bulgaricus Muller, 1979 APPENDIX list of all A: Systematic species and genera currently Genus Tasadia: Tasadia carniolica (Bittner, 1884), as Cancer. the Cancridae. included in each assigned to Species genus are listed after the genus.
I. Family Cancridae, Subfamily Cancrinae APPENDIX B: Key to the Genera ofthe Cancridae,
Genus Cancer sensu stricto: Cancer bellianus Johnson, 1861;
C. borealis Stimpson, 1859; C. fissus Rathbun, 1908; c. 1. Carapace with odd number of frontal spines; posterolateral
fujinaensis Sakumoto, Karasawa, and Takayasu, 1992; c. margin smooth or with oneor two small spines
irroratus Say, 1817; C. johngarthi Carvacho, 1989; (C. Cancrinae and 2
C. Collins and 1'. with evennumber offrontal pagurus , Linnaeus, 1758; parvidens Fraaye, Carapace spines; posterolateral
1991; C. porteri Rathbun, 1930; C. plebejus Poeppig, 1836; margin with numerous coarse granules or spines
C. productus Randall, 1839. Lobocarcininae and 9
2. smooth Genus Anatolikos: Anatolikos japonicus (Ortmann, 1893) new Carapace regions separated by very deep, narrow,
surface of anterolateral comb,, as Cancer; A. itoigawai (Karasawa, 1990)new comb., grooves; regions densely granular; spines
as Cancer (Glebocarcinus); and A. tumifrons (Yokoya, 1933) small, wider than high Platepistoma
2'. weak new comb., as Cancer. Carapace regions separated by grooves or indistinct;
ornamentation variable but not as above; Genus Anisospinos: Anisospinos berglundi new species; carapace region
anterolateral better 3 Anisospinos odosensis (Imaizumi, 1962), new comb., as spines typically developed, longer
Cancer ; and A. wahkiakumensis (Berglund and Goedert, 3. Anterolateral spines long, separated to bases 4
3'. Anterolateral fissured to 1992) new comb., as Cancer (Romaleon). spines predominantly or separated
base and short, or serrate 6 Genus Glebocarcinus ; Glebocarcinus amphioetus (Rathbun, granular,
4. not much wider than long; regions distinct, 1898), as Cancer and G. oregonensis (Rathbun, 1898), as Carapace separated wide, Cancer. by smooth, grooves; regions densely granular on most
Genus Metacarcinus: Metacarcinus anthonyi (Rathbun, 1897) elevated portion Glebocarcinus
4'. much wider than as Cancer; M. danaiNations, 1975, as Cancer (Metacar- Carapace long; regions separated by narrow
M. davidi Cancer ornamentation variable 5 cinus); Nations, 1975, as (Metacarcinus); grooves or indistinct; regional
Anterolateral M. edwardsii (Bell, 1835), as Cancer; M. gracilis (Dana, 5. spines singular, curving anteriorly
Romaleon 1852), as Cancer; M. granti (Rathbun, 1932), as Cancer ;
M. izumoensis Sakumoto, Karasawa, and Takayasu, 1992, 5'. Anterolateral spines mostly paired, curving anteriorly, each
ofdifferent size as Cancer (Metacarcinus); M. jenniferae Nations, 1975, as Anisospinos
6. Front orbits directed forward, distinct Cancer (Metacarcinus); M. magisterDana, 1852),asCancer; weakly projected, spines
M. minutoserratus (Nagao, 1941), as Cancer, M. novaezea- 7
6'. Front frontal landiae (Jacquinot, 1853), as Cancer; M. starri Berglund strongly projected, spines coalesced; orbits
directed and Goedert, 1996, as Cancer (Metacarcinus). anterolaterally 8
* 7. Anterolateral fissured, Genus Notocarcinus: Notocarcinus sulcatus new species. spines predominantly strongly may be base Genus Platepisloma: Platepistoma anaglyptumBalss, 1922;P. separated to posteriorly Cancer T. Anterolateral less fissured guezi (Crosnier, 1976), as Cancer; P. imamurae (Imaizumi, spines distinctly or small, separated
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to serrate Metacarcinus 9'. Frontal with four to six anterolateral bases, terminations granular or margin spines; spines
8. Anterolateral spines mostly paired, terminations sharp, regions sharp, separated to bases, often bifurcate or trifurcate; postero- inflated Anatolikos lateral margin with large granules or spines 10
8'. Anterolateral spines singular, terminations blunt, epigastric 10. Anterolateral spines obviously bifurcate or trifurcate; and hepatic regions sulcate Notocarcinus posterolateral margin with prominent spines Lobocarcinus
9. Frontal margin with six spines; anterolateral spines fissured, 10'. Anterolateral spines singular or weakly bifurcate; with blunt terminations except for ninth which is sharp; posterolateral margin with coarse granules Tasadia posterolateral margin smooth Miocyclus
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