A New Classification of the Xanthoidea Sensu Lato
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Contributions to Zoology, 75 (1/2) 23-73 (2006) A new classifi cation of the Xanthoidea sensu lato (Crustacea: Decapoda: Brachyura) based on phylogenetic analysis and traditional systematics and evaluation of all fossil Xanthoidea sensu lato Hiroaki Karasawa1, Carrie E. Schweitzer2 1Mizunami Fossil Museum, Yamanouchi, Akeyo, Mizunami, Gifu 509-6132, Japan, e-mail: GHA06103@nifty. com; 2Department of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720, USA, e-mail: [email protected] Key words: Crustacea, Decapoda, Brachyura, Xanthoidea, Portunidae, systematics, phylogeny Abstract Family Pilumnidae ............................................................. 47 Family Pseudorhombilidae ............................................... 49 A phylogenetic analysis was conducted including representatives Family Trapeziidae ............................................................. 49 from all recognized extant and extinct families of the Xanthoidea Family Xanthidae ............................................................... 50 sensu lato, resulting in one new family, Hypothalassiidae. Four Superfamily Xanthoidea incertae sedis ............................... 50 xanthoid families are elevated to superfamily status, resulting in Superfamily Eriphioidea ......................................................... 51 Carpilioidea, Pilumnoidoidea, Eriphioidea, Progeryonoidea, and Family Platyxanthidae ....................................................... 52 Goneplacoidea, and numerous subfamilies are elevated to fam- Family Hypothalassiidae ................................................... 52 ily status. The Mathildellidae is moved from the Goneplacidae Family Pseudoziidae .......................................................... 53 to the Portunoidea. Diagnoses for all superfamilies and families Family Eriphiidae ............................................................... 54 discussed herein are provided, embracing characters typically Family Oziidae .................................................................... 54 used by biologists as well as readily fossilized features of the Superfamily Progeryonoidea .................................................. 54 dorsal carapace, sternum, abdomen, and chelipeds. All genera Family Progeryonidae ....................................................... 54 known from the fossil record at one time referred to the Xanthi- Superfamily Goneplacoidea ................................................... 55 dae sensu lato, Xanthoidea sensu lato, or Goneplacidae sensu Family Carinocarcinoididae ............................................. 56 lato were evaluated as to their family level placement and as a Family Chasmocarcinidae ................................................ 56 result, the family-level placement of many of these genera has Family Euryplacidae .......................................................... 57 been changed herein. Balcacarcinus new substitute name, is Family Goneplacidae ......................................................... 57 provided herein for Bittneria Schweitzer and Karasawa, 2004. Family Hexapodidae .......................................................... 58 Family Trogloplacidae ....................................................... 58 Superfamily Goneplacoidea incertae sedis ......................... 58 Superfamily Portunoidea ........................................................ 59 Contents Family Carcineretidae ....................................................... 59 Family Geryonidae ............................................................. 60 Introduction ...................................................................................... 23 Family Mathildellidae ....................................................... 61 Phylogenetic analysis of the Xanthoidea sensu lato ............... 25 Family Portunidae .............................................................. 61 Material and methods .............................................................. 25 The fossil record and diversity of Results ......................................................................................... 27 the Xanthoidea sensu lato ....................................................... 62 Discussion .................................................................................. 27 Acknowledgements ........................................................................ 64 Systematic paleontology: superfamilies and families References ........................................................................................ 65 recognized herein ...................................................................... 41 Superfamily Carpilioidea ........................................................ 42 Family Carpiliidae .............................................................. 42 Introduction Family Palaeoxanthopsidae .............................................. 43 Family Tumidocarcinidae ................................................. 44 Family Zanthopsidae .......................................................... 44 The brachyuran superfamily Xanthoidea MacLeay, Superfamily Pilumnoidoidea .................................................. 45 1838, sensu lato is a large, diverse group in modern Family Pilumnoididae ....................................................... 45 oceans, and it has a reasonably good fossil record. Superfamily Xanthoidea .......................................................... 46 Beurlen (1930) fi rst recognized the superfamily (as Family Domeciidae ............................................................ 46 Family Panopeidae ............................................................. 47 his subtribus) Xanthoidea containing seven families, Genus Balcacarcinus ................................................... 47 Gecarcinidae MacLeay, 1838; Geryonidae Colosi, Downloaded from Brill.com09/30/2021 09:19:56PM via free access 24 Karasawa and Schweitzer – A new classifi cation of the Xanthoidea sensu lato 1923; Goneplacidae MacLeay, 1838; Grapsidae Schweitzer, 2003a; Tumidocarcinidae Schweitzer, MacLeay, 1838; Pinnotheridae De Haan, 1833; 2005a; and Zanthopsidae Vía, 1959. Potamidae Ortmann, 1896; and Xanthidae MacLeay, Recent investigations have resulted in the descrip- 1838 sensu lato. In her classifi cation of brachyurans, tion of new fossil and extant material, and numerous Guinot (1978) included eight families within the studies have revisited fossil and extant material. Xanthoidea: Carpiliidae Ortmann, 1893; Menippidae Thus, revisions to the systematics and classifi cation Ortmann, 1893; Platyxanthidae Guinot, 1977; Xan- of the currently known xanthoid families have been thidae; Panopeidae Ortmann, 1893; Pilumnidae provided for the following families which include Samouelle, 1819; Trapeziidae Miers, 1886; and extant and extinct taxa: Carpiliidae, Palaeoxanthop- Geryonidae. Manning and Holthuis (1981) fi rst sug- sidae, Pseudoziidae, and Zanthopsidae (Schweitzer, gested that the Geryonidae had close affi nities with 2003a); Eriphiidae, Platyxanthidae, and Tumidocar- the Por tunidae, and subsequently, Bowman and cinidae (Schweitzer, 2005a); Domeciidae and Tra- Abele (1982) placed the Geryonidae within the peziidae (Castro et al., 2004; Schweitzer, 2005b); Portunoidea and recognized four families within the Goneplacidae (Karasawa and Kato, 2003a, b); Hexa- Xanthoidea: Goneplacidae, Hexapodidae, Xanthi- podidae (Schweitzer and Feldmann, 2001); Pilum- dae, and Platyxanthidae. nidae (Schweitzer, 2000); and Panopeidae and Subsequently, work on the classifi cation of the Pseu dorhombilidae (Schweitzer and Karasawa, Xanthoidea has continued in earnest. Jamieson (1993) 2004). These studies were performed using cladistic fi rst suggested that the Xanthoidea was paraphylet- analysis based upon adult morphology or using ic based upon spermatological evidence. Coelho and proxy characters of the dorsal carapace. Coelho Filho (1993) considered the Xanthidae Among the various families discussed above and sensu lato to be polyphyletic based upon phenetic assigned to the Xanthoidea sensu lato, the Eumedo- and cladistic analysis and divided it into four fami- nidae has been considered to be one of the sub- lies: Carpiliidae; Xanthidae including the Menip- families within the Pilumnidae (Ng and Clark, 2000; pinae, Platyxanthinae, Xanthinae, and Eucratopsinae; Davie, 2002; Poore, 2004); we concur. We follow Eriphiidae; and Pilumnidae including the Trapeziinae Schweitzer (2005b) in considering the Trapeziidae and Pilumninae. The monophyly of the Xanthoidea and Tetraliidae as a single family, and our analysis was further questioned based upon additional cla- showing genera from each group as sisters supports distic analyses using adult morphology (Von Stern- her decision. berg et al., 1999; Von Sternberg and Cumberlidge, Števcˇic´ (2005) published a major revision of the 2001; Castro et al., 2004), foregut ossicles (Brösing, brachyuran families including both fossil and extant 2002), and molecular data (Schubart et al., 2000; forms, one the few such major works to do so. Among Wetzer et al., 2003). Martin and Davis (2001) in- the Xanthoidea sensu lato, his revision includes cluded as xanthoid families the Carpiliidae; Eume- numerous new subfamilies and tribes, nearly all for donidae Dana, 1853; Goneplacidae; Hexapodidae extant genera. He also raised some subfamilies to Miers, 1886; Menippidae (= our Eriphiidae, see family status. Because our work has concentrated Davie [2002] and Schweitzer