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Re-Evaluation of the Cancridae Latreille, 1802 (Decapoda: Brachyura) Including Three New Genera and Three New Species 4 Carrie E

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Re-Evaluation of the Cancridae Latreille, 1802 (Decapoda: Brachyura) Including Three New Genera and Three New Species 4 Carrie E WW L^^ii»~^i • **«^«' s^f J-CLI//^/*IP / +' IU ! U~~> O' S~*Af/^i Contributions to Zoology, 69 (4) 223-250 (2000) SPB Academic Publishing hv. The Hague Re-evaluation of the Cancridae Latreille, 1802 (Decapoda: Brachyura) including three new genera and three new species 4 Carrie E. Schweitzer & Rodney M. Feldmann Department of Geology, Kent State University, Kent, Ohio 44242 U.S.A. E-mail: [email protected]. edit and [email protected] Keywords: Decapoda, Brachyura, Cancridae, Tertiary, paleobiogeography, Tethys Abstract Metacarcinus A. Milne Edwards, 1862 235 Metacarcinus goederti new species 236 New fossils referrable to the Cancridae Latreille, 1802 extend Notocarcinus new genus 239 the known stratigraphic range of the family into the middle Eocene Notocarcinus sulcatus new species 240 and the geographic range into South America. Each genus within Platepistoma Rathbun, 1906 241 the family has been reevaluated within the context of the new Romaleon Gistl, 1848 242 material. A suite of diagnostic characters for each cancrid ge­ Subfamily Lobocarcininae Beurlen, 1930 243 nus makes it possible to assign both extant and fossil speci­ Lobocarcinus Reuss, 1867 243 mens to genera and the two cancrid subfamilies, the Cancrinae Miocyclus Miiller, 1979 244 Latreille, 1802, and Lobocarcininae Beurlen, 1930, based solely Tasadia Miiller in Janssen and Miiller, 1984 244 upon dorsal carapace morphology. Cheliped morphology is useful Discussion 245 in assigning genera to the family but is significantly less useful Acknowledgements 246 at the subfamily and generic level. Each of the four subgenera References 246 sensu Nations (1975), Cancer Linnaeus, 1758, Glebocarcinus Appendix A 249 Nations, 1975, Metacarcinus A. Milne Edwards, 1862, and Appendix B 249 Romaleon Gistl, 1848, are elevated to full generic status. Addi­ tionally, three new genera and three new species accommodate the new, as well as some previously described taxa, and include Introduction Anatolikos new genus, Anisospinos berglundi new genus and species, and Notocarcinus sulcatus new genus and species and several new combinations. Recognition of new genera and The Cancridae Latreille, 1802, has heretofore con­ reassignment of several species within the Cancrinae indicates sisted of two extant genera known from the fossil that that subfamily may have arisen in the southern hemisphere, record, Cancer sensu lato and Platepistoma Rath- contrary to the previous interpretation of the subfamily as a primarily North Pacific or Tethyan group. The Lobocarcininae bun, 1906, and several exclusively fossil genera was primarily a Tethyan group. including Lobocarcinus Reuss, 1867; Microdium Reuss, 1867; Miocyclus, Miiller, 1979; Parapirimela Van Straelen, 1937; and Tasadia Miiller in Janssen Contents and Miiller, 1984. Cancer sensu lato was subdivided into four subgenera by Nations (1975) including Introduction 223 Cancer, Glebocarcinus, Metacarcinus, and Roma­ Systematic Paleontology 224 leon. All of these genera and subgenera were de­ Family Cancridae Latreille, 1802 224 fined primarily upon the number, size, and shape Subfamily Cancrinae, Latreille, 1802 226 of the frontal spines; the number, size, ornament, Cancer sensu stricto Linnaeus, 1758 227 and shape of the anterolateral spines; the ornament Anatolikos new genus 228 Anisospinos new genus 229 of the posterolateral margin; the shape, develop­ Anisospinos berglundi new species 231 ment, and ornament of carapace regions, and the Glebocarcinus Nations, 1975 234 shape and ornament of the chelae. 224 C.E. Schweitzer & R.M. Feldmann - Re-evaluation ofCancridae The discovery of new fossil material from Alaska Systematic Paleontology and Washington, USA, and southern Argentina requires a re-evaluation of the family. Within this Order Decapoda Latreille, 1802 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. Neither Parapirimela nor Branchio- than eight; posterolateral margin entire, rimmed lambrus are verifiable members of the Cancridae or spined; carapace regions ranging from indis­ as discussed below. Additionally, the morphology tinct 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 cantly less useful in assigning taxa to genera, be­ shape; sternites 3-5 often fused in males. "Anten- cause of the variability in chela morphology. nules folded lengthwise. Antennal flagella present, The evolutionary and biogeographic history of short, more or less hairy. Third maxillipeds usu­ the group must be reinterpreted within the context ally overlapping endostome" (Rathbun, 1930, p. of the new fossil material. The discovery of Noto­ 176). "Carapace broadly oval or hexagonal. Last carcinus sulcatus new genus and species from pair of legs not adapted for swimming" (Williams, middle Eocene rocks of Argentina constitutes the 1984, p. 351). earliest known occurrence of the Cancrinae and also the first known fossil occurrence of the Can­ Remarks. - This diagnosis (and all the diagnoses cridae in South America. Anisospinos berglundi to follow) focus on characters that are typically is the only other Eocene 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 Washington. It is possible that the Cancrinae arose ing the family is a high degree of 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 Oceans, where it is currently well-estab­ members of the family possess mani of the first 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 1979; Carvacho, 1989) or Tethyan (Newman, 1991) key to all genera herein included within the family distribution. Most of the genera assigned to the (Appendix B; Fig. 1), employs these characteristics Cancrinae in this report, including Anatolikos new to distinguish taxa. Miiller (1984) placed the genus genus, Anisospinos new genus, Cancer sensu stricto, Microdium, within the Cancridae, and suggested Platepistoma, and Romaleon, apparently originated that it may be closely allied to or synonymous with in the North Pacific region based upon known Cancer s. I. However, the holotype for the sole occurrences. Assignment of the European genera species, M. nodulosum, has been lost (Miiller, 1984), Miocyclus and Tasadia to the Lobocarcininae sup­ so confirmation of the status of that genus is im­ ports the primarily Tethyan distribution of that possible. According to Miiller (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 Contributions to Zoology, 69 (4) - 2000 225 mm I TASADIA 1. LOBOCARCINUS 4\ MIOCYCLUS l< I? S .NOTOCARC/NUS i 1 cc l< 18 ANA TOLIKOS !S \ i METACARCINUS h- CANCER / I II I AN I SOSPIN OS II I Mlui -Hi M,< ROMALEON *>' \l !< o «/ I GLEBOCARCINUS PLATEPISTOMA > CENOZOIC Fig. 1. Illustrated key to the subfamilies and genera of the Cancridae showing the geologic range of each genus. The branching structure of the key does not reflect phylogeny; each node represents a couplet in the dichotomous key (Appendix B). 226 C.E. Schweitzer & R.M. Feldmann - Re-evaluation of Cancridae Parapirimela within the Cancridae. The sole spe­ Remarks. - The Cancrinae, until now, has been a cies, P. angolensis van Straelen, 1937, was founded small subfamily comprised of only a few genera, upon a fragment of dorsal carapace and cheliped. Cancer s. I. and Platepistoma as well as Para­ The anterolateral margin was described as possess­ pirimela, Microdium, and Branchiolambrus, which ing four anterolateral spines, the last of which is have been discussed above. This report introduces bifid; a front with five spines including the poorly three new genera to the subfamily; in addition, the developed inner orbital spines; and a carapace that four subgenera of Cancer s. I., Cancer s. s., Gle­ is wider than long. None of these characteristics bocarcinus, Metacarcinus, and Romaleon, are can be seen on the illustrations of this taxon. The herein elevated to generic status, bringing the holotype is described as housed in a private col­ number of included genera to eight. Diagnoses and lection and may, therefore, be lost. Thus, it is im­ included species for each genus are provided be­ possible to confirm placement
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