Checklist of fossil decapod crustaceans from tropical America. Part I: Anomura and BrachyuraAbstract
Our knowledge of fossil crustaceans from the tropics has increased considerably during recent decades, thanks to novel findings and the re-examination of museum specimens. However, several previous records have been misidentified, numerous museum specimens have never been reported, and many new discoveries are yet to be published. Here, we present a detailed, up-to-date, and revised checklist for every marine, terrestrial, or freshwater fossil decapod crustacean occurrence from tropical America known to us, including their age, geographic occurrences, and related literature. We recognize the occurrence of at least 32 superfamilies, 69 families, 190 genera, and 415 species of brachyurans (‘true’ crabs), and anomurans (‘false’ crabs, hermit crabs, squat lobsters, and allies), several of them previously unknown. The checklist comprises records from three main geographic regions: 1) northern South America (Bolivia, Brazil, Chile, Colombia, Ecuador, Peru, Venezuela); 2) Central America and southern North America (Belize, Costa Rica, Honduras, Panama, Mexico, southern and central Florida); and 3) the Caribbean Islands + Bermuda (Anguilla, Antigua, Aruba, Bahamas, Barbados, Bermuda, Bonaire, Cuba, Curaçao, Dominican Republic, The Grenadines, Haiti, Jamaica, Puerto Rico, Saint Bartélemy, Saint Martin, Trinidad). Previous findings, new occurrences, and the revised systematic placement for several problematic/misidentified records, indicate that the fossil record of anomurans and brachyurans in tropical America is more diverse than previously envisioned, with a considerable degree of endemism at the genus- and species-levels.
Key words
Cenozoic; Central America; Mesozoic; Neotropics; North America; South America
Table of contents
Introduction .................................................................... 2
Institutional abbreviations ........................................... 7
Annotated checklist ....................................................... 7
Northern South America .............................................. 7
Bolivia ......................................................................... 7
Brazil ........................................................................... 8
Chile ......................................................................... 15
Colombia ..................................................................... 18
Ecuador .................................................................... 22
Peru ........................................................................... 23
Venezuela ................................................................. 24
Central America and southern North America ..... 29
Belize ........................................................................ 29
Costa Rica ................................................................ 29
Honduras .................................................................. 32
Panama ..................................................................... 32
Mexico ........................................................................ 38
Southern and central Florida, USA .................... 47
Caribbean Islands & Bermuda ................................... 51
Anguilla ..................................................................... 51
Antigua ..................................................................... 52
Aruba ........................................................................ 54
Bahamas ................................................................... 54
Barbados .................................................................. 54
Bermuda ................................................................... 55
Bonaire ..................................................................... 56
Cuba .......................................................................... 56
Curaçao .................................................................... 58
Dominican Republic ............................................. 59
The Grenadines ...................................................... 61
Haiti .......................................................................... 61
Jamaica ..................................................................... 62
Puerto Rico ............................................................. 68
Saint Barthélemy (St. Barts) ................................ 69
Saint Martin ............................................................ 69
Trinidad ................................................................... 69
Final Remarks .............................................................. 70
Acknowledgments ....................................................... 71
References ...................................................................... 72
Introduction
New paleontological discoveries made mostly during the last two decades have dramatically expanded our understanding of the crustacean fossil record from the New World’s tropics. Recent works on fossil and extant anomurans and brachyurans (Hendrickx, 1995; Magalhães, 2003; Ng et al., 2008; De Grave et al., 2009; Boyko and McLaughlin, 2010; McLaughlin et al., 2010; Osawa and McLaughlin, 2010; Schweitzer et al., 2010; Jagt et al., 2015; among many others) have been pivotal to the synthesis of the current state-of-knowledge of marine, terrestrial, and freshwater fossil decapods in tropical America: one of the most diverse regions on Earth. These new discoveries, together with novel phylogenetic hypotheses, and re-examinations of several previous findings, call for an urgent revision of the crustacean fossil record from tropical America, and are the motivation behind the present checklist. In this work, we focus on the brachyuran and anomuran fossil decapods from the tropical Western Hemisphere and the need to put them into updated temporal, geographic, and systematic contexts.
The tropics are defined as the regions surrounding the Equator, extending roughly from the Tropic of Cancer (~23.3˚N) to the Tropic of Capricorn (~23.3˚S). In addition to the tropical occurrences of fossil anomurans and brachyurans, we have included records from subtropical areas ranging in latitude from ~30˚N to ~30˚S (Fig. 1). Since the Neotropical region engulfs most of South America, here we restrict the South American fossil records to those north of 30˚S latitude. The Caribbean, for the purpose of our work, is considered to be the non-continental land surrounded by the Caribbean Sea, and mostly positioned on the Caribbean tectonic plate. The fossil occurrences have been grouped into three main geographic regions: 1) northern South America, with records from Bolivia, Brazil, Chile, Colombia, Ecuador, Peru, and Venezuela; 2) Central America and southern North America, with records from Belize, Costa Rica, Honduras, central and southern Florida, Mexico, and Panama; and 3) the Caribbean Islands + Bermuda, with records from Anguilla, Antigua, Aruba, Bahamas, Barbados, Bermuda, Bonaire, Cuba, Curaçao, Dominican Republic, the Grenadines, Haiti, Jamaica, Puerto Rico, Saint Bartélemy, Saint Martin, and Trinidad (Fig. 1). To the best of our knowledge, these are the tropical American countries, states, territories, or islands from where fossil anomurans and brachyurans have been either reported in the literature, or found as unpublished material in museum collections (e.g., USNM, MNHN) and reported here. Other tropical American countries, territories, or islands not listed or mentioned lack known brachyuran or anomuran fossils.
Location map of the tropical and subtropical American countries, states, territories, or islands where fossil anomurans and brachyurans have been discovered/reported to date. A, northern South America (light gray) with records from Bolivia, Brazil, Chile, Colombia, Ecuador, Peru, and Venezuela; continental Central America and southern North America (intermediate gray), with records from Belize, Costa Rica, Honduras, Panama, Mexico, and southern and central Florida; B, Caribbean Islands and Bermuda (dark gray), with records from Anguilla, Antigua, Aruba, Bahamas, Barbados, Bermuda, Bonaire, Cuba, Curaçao, Dominican Republic, the Grenadines, Haiti, Jamaica, Puerto Rico, Saint Bartélemy, Saint Martin, and Trinidad; C, close-up of the ABC islands north of Venezuela: Aruba, Bonaire, and Curaçao; D, count of confirmed (grey bars) and indeterminate (black bars) number of families, genera, and species of fossil anomurans and brachyurans for each of the countries, states, territories, or islands included in this study.
This work provides not only an updated and detailed list of fossil anomurans and brachyurans from tropical America, but also includes several new records and a re-examination of the systematic placement of problematic taxa. Although some genera such as Lobonotus A. Milne-Edwards, 1863, AraripecarcinusMartins-Neto, 1987, TepexicarcinusFeldmann, Vega, Applegate and Bishop, 1998b, PrehepatusRathbun, 1935b, and RoemerusBishop, 1983b, still have unclear systematic affinities due to convergence, incompleteness of their carapaces, and/or poor preservation (e.g., Bishop, 1985; Vega et al., 1995b; 2005; 2006a; Schweitzer et al., 2006b; Jagt et al., 2010; 2014; Ossó et al., 2014; Luque, 2015a; 2015b), the systematic position of most other families, genera, and species here included have been verified by us and are, to the best of our knowledge, as accurate and updated as possible. As a result, we recognize the occurrence in the tropical Americas of at least 32 superfamilies, 69 families, 190 genera, and 415 spp. of brachyurans (‘true’ crabs) and anomurans (‘false’ crabs, hermit crabs, squat lobsters, and allies) (Chart. 1). New records include the first occurrence of early Cretaceous pagurid anomurans from Colombia, the crabs CostacoplumaCollins and Morris, 1975, QuasilaeviraninaTucker, 1998, and PalaeoxanthopsisBeurlen, 1958 from the late Cretaceous of Colombia; EuphylaxStimpson, 1862 from the Eocene of Peru; ParaeuphylaxVarela and Schweitzer, 2011 from the Miocene of Venezuela; PortunusWeber, 1795 from the Miocene of Colombia; JohngarthiaTürkay, 1970 from the Pleistocene of Brazil; FalconoplaxVan Straelen, 1933 and PinnixaWhite, 1846 from the Miocene of Panama; IxaLeach, 1817 from the Cenozoic Chile; and some eubrachyurans from the Cenomanian of Bolivia and the Paleocene of Colombia.
Geographic (left) and stratigraphic (right) distribution of 32 superfamilies and 69 families of anomurans and brachyurans with fossil records in tropical America summarized in this work. Colors for countries/regions as in Fig. 1. Colors for families and superfamilies as in Fig. 2. Fossil occurrences for a given family and country are marked with ‘X’. Solid horizontal bars indicate the known chronologic and stratigraphic ages for a given family, dotted lines indicate unknown but expected occurrences, and outlined white bars indicate taxa with dubious or indeterminate familial placement. Taxa marked with a dagger (†) only known from fossils. SC Florida = South and Central Florida. Numerical ages given in millions of years (Ma). Geologic time abbreviations: AL = Albian, AP = Aptian, BA = Barremian, BE = Berriasian, CA = Campanian, CE = Cenomanian, CO = Coniacian, E = early, HA = Hauterivian, Holo = Holocene, L = late, MA = Maastrichtian, M = middle, Oligo = Oligocene, Paleo = Paleocene, Pleisto = Pleistocene, Plio = Pliocene, SA = Santonian, TU = Turonian, VA = Valanginian. Geologic time slices not to scale.
Although the phylogenetic position of most brachyuran families is relatively well resolved, there are still discrepancies regarding the classification and subsequent naming of the principal brachyuran ranks above superfamily- and below infraorder-levels, i.e., sections and subsections. The International Code of Zoological Nomenclature does not regulate the nomenclature for taxa in ranks between superfamily and infraorder, which has led to at least two different phylogenetic hypotheses: a) a monophyletic Podotremata (e.g., Guinot et al., 2013; Davie et al., 2015; Jagt et al., 2015), and b) a paraphyletic podotremous grade (e.g., Ahyong et al., 2007; De Grave et al., 2009; Scholtz and McLay, 2009; Karasawa et al., 2011; Tsang et al., 2014). Since the aim of the present work is to provide a revised and updated list of fossil decapod crustaceans from tropical America and not to discuss the phylogenetic relationships among higher brachyuran taxa between the superfamily and infraorder ranks, we have grouped the taxa under superfamilies, first based on their approximate phylogenetic position (e.g., typical podotremous superfamilies are listed before eubrachyuran superfamilies) (Fig. 2), and second, alphabetically within superfamilies. The schematic phylogenetic relationships among the anomuran and brachyuran superfamilies and sections/subsections listed in this work are partially based on the works of Bracken-Grissom et al. (2013) for Anomura (white box), Karasawa et al. (2011) for podotremous Brachyura (colored boxes), and Tsang et al. (2014) for eubrachyuran Brachyura (grey box) (Fig. 2; Chart 1). In this checklist, one dagger (†) indicates taxa that are exclusively known from fossils, and no dagger denotes extant taxa with known fossil record. Extant-only species, genera, families and superfamilies are not listed here.
Schematic phylogenetic relationships among the superfamilies of Anomura and Brachyura listed in this work. The general topology and colored boxes reflect current phylogenetic scenarios based on molecular and morphological data, partially following the works of Bracken-Grissom et al. (2013) for Anomura (white box), Karasawa et al. (2011) for podotremous Brachyura (colored boxes), and Tsang et al. (2014) for some eubrachyuran Brachyura (grey box). The order in which superfamilies are listed in this figure, from top to bottom, is the same followed through the checklist and in Chart 1, whereas families, genera, and species within a given superfamily are listed alphabetically. Superfamilies with a dagger (†) are only known from fossils.
INSTITUTIONAL ABBREVIATIONS
CPC = Colección Paleontológica de Coahuila, Museo del Desierto, Saltillo, Coahuila, Mexico.
IGM = Colección Nacional de Paleontología, Instituto de Geología, UNAM, Mexico D.F., Mexico.
IGM p = Colecciones Paleontológicas Museo José Royo y Gómez, Servicio Geológico Colombiano, Bogotá, Colombia (formerly INGEOMINAS)
IHNFG = Colección Paleontológica de Chiapas, Museo Eliseo Palacios, Tuxtla Gutiérrez, Chiapas, Mexico.
LPURCA = Laboratório de Paleontologia da Universidade Regional do Cariri.
MNHN = Muséum national d’Histoire naturelle, Paris, France.
MNRJ = Museu Nacional, Universidade Federal do Rio de Janeiro.
MUN STRI = Mapuka Museum of Universidad del Norte, Barranquilla, Colombia.
MUZ = Museo de Múzquiz, Múzquiz, Coahuila, Mexico.
UF = Invertebrate Paleontology Division, Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA.
USNM = United States National Museum, Smithsonian Institution, Washington, D.C., USA.
Annotated checklist
Order DECAPODA Latreille, 1802
Clade MEIURA Saint Laurent, 1980
Northern South America
BOLIVIA
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Diogenidae incertae sedis ENT#091;Cenomanian, PotosíENT#093; (Fig. 3A-H) ENT#091;Notes 1, 2ENT#093;.
Fossil Anomura and Brachyura from the Cretaceous of Bolivia, South America. A-H, Anomura: Paguroidea: ?Diogenidae, Cenomanian of Potosí; A, B, MNHN-F.A57970, left cheliped in outer (A) and inner (B) views; C, D, MNHN-F.A57972, left cheliped in outer (C) and inner (D) views; E-H, MNHN-F.A57971, left chelipeds in outer (E, G) and inner (F, H) views. I-L, Brachyura: Eubrachyura, of Potosí; I, J, eubrachyuran indet., MNHN-F.A57973, in dorsal (I) and ventral (J) view. K-L, eubrachyuran indet., MNHN-F.A57966, in dorsal (K) and ventral (L) view. Photos courtesy of Jocelyn Falconnet, Peter Massicard, and Sylvain Charbonnier (MNHN). All specimens dry, uncoated.
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily ?CARPILIOIDEA Ortmann, 1893
?Tumidocarcinidae Schweitzer, 2005a
Dynomenopsis † Secretan, 1972
Dynomenopsis branisai † Secretan, 1972 (type) ENT#091;Cenomanian, PotosíENT#093; (Schweitzer and Feldmann, 2012) ENT#091;Note 2ENT#093;.
EUBRACHYURA incertae sedis
Eubrachyuran 1 (carapace) ENT#091;Cenomanian, PotosíENT#093; (Fig. 3I, J) ENT#091;Note 2ENT#093;.
Eubrachyuran 2 (carapace) ENT#091;Cenomanian, PotosíENT#093; (Fig. 3K, L) ENT#091;Note 2ENT#093;.
Eubrachyuran indet. (claw fragments) ENT#091;Cenomanian, PotosíENT#093; (Fig. 4A-J) ENT#091;Note 2ENT#093;.
?Eubrachyuran indet. (claw fragments) ENT#091;Miocene, Santa CruzENT#093; (Fig. 4K-N)
Fossil Brachyura from the Cretaceous and Neogene of Bolivia, South America. A-J, eubrachyuran chelipeds associated with crab carapace MNHN-F.A57966 (see Fig. 3K, L) from the Cenomanian of Potosí; A-H, MNHN-F.A57968, several right chelipeds in outer (A, C, E, G) and inner (B, D, F, H) views; I, J, MNHN-F.A57967, a large right cheliped in outer (I) and inner (J) view. K-N, eubrachyuran chelipeds from the Miocene of Santa Cruz; K, MNHN-F.A57963, dactylus; L, MNHN-F.A57961, dactylus; M, MNHN-F.A57962, dactylus; N, MNHN-F.A57964, fragment of pollex. Photos courtesy of Jocelyn Falconnet, Peter Massicard, and Sylvain Charbonnier (MNHN). All specimens dry, uncoated.
Note 1. To date, the only known fossil decapod records from Bolivia are those reported by Secretan (1972), which include some indeterminate decapod cheliped remains, and the brachyuran Dynomenopsis branisai, all from the Cenomanian Miraflores Formation of the Puca Group in Esquena. A parcel of decapod remains deposited at the Paleontological Collections of the Muséum National d’Histoire Naturelle in Paris (MNHN), includes several paguroid chelipeds and many brachyuran carapace and cheliped remains (Figs. 3, 4). The paguroid remains consist mainly of major left chelipeds, which are characteristic of the family Diogenidae (including Coenobitinae), and reminiscent of the ‘indeterminate decapod remains’ reported by Secretan (1972, text fig. 1, plates II and III). Herein, we tentatively assign Secretan’s specimens and the new material to Diogenidae incertae sedis (Fig. 3A-H), until further studies permit a more accurate systematic placement.
Note 2. To our knowledge, Dynomenopsis branisai, from the Cenomanian of Esquena, is the only known fossil brachyuran from Bolivia to date. As suggested by Schweitzer and Feldmann (2012), the affiliation of Dynomenopsis with Tumidocarcinidae, or even Carpilioidea, can only be corroborated as more material becomes available. Here, we maintain it within Tumidocarcinidae albeit with uncertainty. The occurrence of two additional carapaces from apparently different taxa (Fig. 3I, L), increase the number of fossil brachyurans known from Bolivia to three. The labels and metadata associated with these specimens indicate that they were collected in Cenomanian rocks of Esquena. Several of the chelipeds and cheliped fragments here illustrated (Fig. 4A-J) are associated with one of the carapaces (Fig. 3K, L), and are strongly reminiscent of the chelipeds seen among several durophagous eubrachyurans (J. Luque, pers. obs.).
BRAZIL
Infraorder ANOMURA MacLeay, 1838
Superfamily GALATHEOIDEA Samouelle, 1819
Munidopsidae Ortmann, 1898
Brazilomunida † Martins-Neto, 2001
Brazilomunida brasiliensis † (Beurlen, 1965, as Galatheites brasiiensis) (type) ENT#091;Albian, SergipeENT#093;
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Dardanus Paul’son, 1875
Dardanus spp. ENT#091;Paleocene, PernambucoENT#093; (inTávora et al., 2005) ENT#091;Note 1ENT#093;
Infraorder BRACHYURA Latreille, 1802
Superfamily DROMIOIDEA De Haan, 1833
Dynomenidae Ortmann, 1892
Maurimia † Martins-Neto, 2001
Maurimia sergipensis † (Beurlen, 1965, as ?Cyclothyreus sergipensis) (type) (as Distefania sergipensis inSchweitzer et al., 2010) ENT#091;Albian, SergipeENT#093;
Superfamily NECROCARCINOIDEA † Förster, 1968
Necrocarcinoidea incertae sedis
Araripecarcinus † Martins-Neto, 1987
Araripecarcinus ferreirai † Martins-Neto, 1987 (type) ENT#091;early Albian, CearáENT#093; (Luque, 2015b) ENT#091;Note 2ENT#093;.
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides spp. ENT#091;Paleocene, PernambucoENT#093; (Távora et al., 2016) ENT#091;Note 3ENT#093;.
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatella Smith, 1869a
Hepatella amazonica † Beurlen, 1958a ENT#091;early Miocene, ParáENT#093;.
Calappidae De Haan, 1833
Calappidae incertae sedis ENT#091;early Miocene, ParáENT#093; ENT#091;Note 4ENT#093;.
Acanthocarpus Stimpson, 1871
Acanthocarpus obscurus † (Rathbun, 1918, as Mursia obscura) ENT#091;early Miocene, ParáENT#093; (inBeurlen, 1958a) (Fig. 5A).
Fossil Eubrachyura from the Cretaceous of Brazil, South America. A, B, Calappoidea: Calappidae: A, Acanthocarpus obscurus (Rathbun, 1918) MNRJ 4583-I, carapace, dorsal view, early Miocene of Pará; B, Calappa circularis (Beurlen, 1958a), syntype, MNRJ 4619-I, carapace, dorsal view, Miocene of Pará. C, D, Portunoidea: Portunidae: Callinectes paraensis Beurlen, 1958a, male, holotype, MNRJ 4585-I, early Miocene of Pará, dorsal (C) and ventral (D) views. E, F, Macropipidae: Ophtalmoplax brasiliana (Maury, 1930), male, MNRJ 4581-I, Maastrichtian of Paraíba, dorsal (E) and ventral (F) views. G, H, Retroplumoidea: Retroplumidae: Costacopluma nordestina Feldmann and Martins Neto, 1995, male, LPURCA specimen uncatalogued, Paleocene of Pernambuco, dorsal (G) and ventral (H) views. Photos by William Santana. All specimens dry, uncoated.
?Calappa Weber, 1795
?Calappa sp. aff. C. zurcheri † Bouvier, 1899 ENT#091;as Calappilia brooksi inTávora et al., 2005, and Calappilia inRumsey et al., 2016) ENT#091;early Miocene, ParáENT#093; (Vega et al., 2009).
Calappilia † A. Milne-Edwards, 1873
Calappilia circularis † (Beurlen, 1958a, as Calappa circularis) ENT#091;Miocene, ParáENT#093; (Fig. 5B).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae incertae sedis
Typilobus † Stoliczka, 1871
Typilobus unispinatus † Martins-Neto, 2001 ENT#091;Miocene, ParáENT#093;. (as Randallia sp. in Beurlen, 1958a)
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Parthenope Weber, 1795
Parthenope trituberculata † Beurlen, 1958a ENT#091;Miocene, ParáENT#093;.
Superfamily CANCROIDEA Latreille, 1802
Cancridae Latreille, 1802
Cancrinae Latreille, 1802
Cyclocancer † Beurlen, 1958a
Cyclocancer tuberculatus † Beurlen, 1958a (type) ENT#091;early Miocene, ParáENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Macropipidae Stephenson and Campbell, 1960
Ophthalmoplax † Rathbun, 1935b
Ophthalmoplax brasiliana † (Maury, 1930, as Zanthopsis brasiliana) ENT#091; Campanian-Maastrichtian, ParaíbaENT#093; (Pralon et al. 2017) (Fig. 5E, F).
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes tajinensis † Vega, Feldmann, Villalobos-Hiriart and Gío-Argíez, 1999 (sensuTávora et al., 2002) ENT#091;early Miocene, ParáENT#093; ENT#091;Note 5ENT#093;.
Scylla De Haan, 1833
Scylla costata † Rathbun, 1919 ENT#091;Miocene, ParáENT#093; (reported in Beurlen, 1958a, and Távora et al., 2002).
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphyplax septendentatus † Beurlen, 1958a ENT#091; early Miocene, ParáENT#093;.
Portuninae Rafinesque, 1815
Achelous De Haan, 1833
Achelous spinimanus (Latreille, 1819) (sensuTávora et al., 2002) ENT#091;early Miocene, ParáENT#093; ENT#091;Note 5ENT#093;.
Arenaeus Dana, 1851d
Arenaeus cribrarius (Lamarck, 1818) ENT#091;early Miocene, ParáENT#093;.
Callinectes Stimpson, 1862
Callinectes paraensis † Beurlen, 1958a ENT#091;early Miocene, ParáENT#093; (Fig. 5C, D).
Callinectes reticulatus † Rathbun, 1918 (claws only) ENT#091;early Miocene, ParáENT#093;.
Portunus Weber, 1795
Portunus atecuicitlis † Vega, Feldmann, Villalobos-Hiriart and Gío-Argíez, 1999 (sensuTávora et al., 2002) ENT#091;early Miocene, ParáENT#093;.
Portunus haitensis † Rathbun, 1923a ENT#091;Miocene, ParáENT#093; (sensuTávora et al., 2002).
Portunus pirabaensis † Martins-Neto, 2001 ENT#091;Miocene, ParáENT#093;.
Portunus spp. ENT#091;early Miocene, ParáENT#093; (cited in Beurlen, 1958a, as Neptunus sp., and in Távora and Dias, 2016, as Portunus oblongus, and erroneously as Palaeopinnixa perornata) ENT#091;Note 6ENT#093;.
Superfamily RETROPLUMOIDEA Gill, 1894
Retroplumidae Gill, 1894
?Archaeopus † Rathbun, 1908
?Archaeopus rathbunae † Beurlen, 1965 ENT#091;Albian, SergipeENT#093; ENT#091;Note 7ENT#093;.
Costacopluma † Collins and Morris, 1975
Costacopluma nordestina † Feldmann and Martins Neto, 1995 ENT#091;Paleocene, PernambucoENT#093; (Fig. 5G, H).
Superfamily TRICHODACTYLOIDEA H. Milne Edwards, 1853
Trichodactylidae H. Milne Edwards, 1853
Trichodactylidae spp. indet. ENT#091;late Miocene, AmazonasENT#093; (Klaus et al., 2017).
Superfamily CARPILIOIDEA Ortmann, 1893
Palaeoxanthopsidae † Schweitzer, 2003
Palaeoxanthopsis † Beurlen, 1958b
Palaeoxanthopsis cretacea † (Rathbun, 1902) (type) ENT#091;Maastrichtian, ParaibaENT#093;.
Tumidocarcinidae † Schweitzer, 2005a
Paratumidocarcinus † Martins-Neto, 2001
Paratumidocarcinus marajoarus † Martins-Neto, 2001 (type) ENT#091;Miocene, ParáENT#093;
Lobonotus † A. Milne-Edwards, 1863
Lobonotus sturgeoni † (Feldmann, Bice, Schweitzer, Salva and Pickford, 1998a) ENT#091;Paleocene, PernambucoENT#093; (in Távora et al., 2005).
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Panopeus H. Milne Edwards, 1834
Panopeus capanemaensis † Martins-Neto, 2001 ENT#091;Miocene, ParáENT#093;
Panopeus sp. ENT#091;early Miocene, ParáENT#093; (in Brito, 1971)
Tetraxanthus Rathbun, 1898
Tetraxanthus rathbunaeChace, 1939 (sensuTávora et al., 2002) ENT#091;Miocene, ParáENT#093;.
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Johngarthia Türkay, 1970
Johngarthia lagostoma H. Milne Edwards, 1837 (as Gecarcinus lagostoma) ENT#091;Pleistocene, PernambucoENT#093; (Fig. 6) ENT#091;Note 8ENT#093;.
Fossil Eubrachyura from the Pleistocene of Brazil, South America. A-L, Grapsoidea: Gecarcinidae: Johngarthia lagostoma H. Milne Edwards, 1837, USNM 618300, Pernambuco; A, B, articulated minor cheliped; C-H, isolated dactyli of major cheliped; I, L, isolated/fragmented cheliped pollices; K, L, isolated/fragmented cheliped pollices. Photos by Javier Luque. All specimens dry, uncoated.
Sesarmidae Dana, 1851c
Sesarma Say, 1817
Sesarma paraensis † Beurlen, 1958a ENT#091;Miocene, ParáENT#093;.
Superfamily OCYPODOIDEA Rafinesque, 1815
Ocypodidae Rafinesque, 1815
Uca Leach, 1814
Uca antiqua † Brito, 1972 (junior synonym U. inaciobritoi † Martins-Neto, 2001) ENT#091;Miocene, ParáENT#093; ENT#091;Note 9ENT#093;.
BRACHYURA incertae sedis
Brachyuran zoea larvae indet. ENT#091;early Albian, CearáENT#093; (Maisey and Carvalho, 1995; Luque, 2015b) ENT#091;Note 2ENT#093;.
Piloslambrus guerini
Note 1. Távora et al. (2005) reported the occurrence of the extant hermit crabs Dardanus fucosusBiffar and Provenzano Jr, 1972, and Dardanus insignis (Saussure, 1858), from the Paleocene Maria Farinha Formation, State of Pernambuco, based on a handful of fragmentary cheliped remains. However, these ~60 million year old fossil specimens do not seem conspecific with D. fucosus or D. insignis, and their systematic affinities need to be re-examined.
Note 2. The holotype and sole specimen of Araripecarcinus ferreirai is a ventral molt, which complicates its systematic placement. Considered by Martins-Neto (1987) as a dorsal carapace of a portunid crab, Guinot and Breton (2006) recognized its superficial resemblance to raninoidan crabs. Karasawa et al. (2008) corroborated the raninoidan affinities. Luque (2015b) re-described and re-illustrated the type specimen of Araripecarcinus, and based on taxonomic and cladistics approaches, indicated that it may be closer to the necrocarcinoid-like clade of raninoidans. Furthermore, its geographic range and age also matches that of other early Cretaceous necrocarcinoids from tropical South America (e.g., Vega et al., 2010; Luque et al., 2012; Karasawa et al., 2014; Luque, 2014). Although Araripecarcinus might be closer to Necrocarcinidae or Orithopsidae, its systematic affinities remain unclear (Luque, 2015b). The only other brachyuran remains known from the Romualdo Formation are a few zoea larvae preserved as fish stomach contents (Maisey, 1994; Maisey and Carvalho, 1995; Luque, 2015b).
Note 3. Távora et al. (2016) reported on a couple of specimens of Raninoides from the Paleocene Maria Farinha Formation that were assigned to two known fossil species from the Eocene and Oligocene of USA: Raninoides fulgidusRathbun, 1926, and Raninoides lewisanaRathbun, 1926. However, the authors did not provide any discussion or remarks justifying their systematic placement. Several species of fossil Raninoides are known from Paleogene rocks worldwide, and most of them are difficult to tell apart from one another. Thus, based solely on the illustrations of Távora et al. (2016) we cannot confirm the specific affinities of the material, and for this reason, we consider them as Raninoides sp. indet. until detailed comparison is made or better material becomes available.
Note 4. Távora et al. (2002, fig. 8) reported a presumably indeterminate species of Callinectes (Portunidae) from the Miocene of Pará. Based solely on their original illustration, we conclude that the specimen does not belong to Callinectes or even Portunidae, but rather represents a dorsal carapace akin to Calappidae.
Note 5. Távora et al. (2002) reported the occurrence of the extinct Necronectes tajinensis and Scylla costata, and the extant portunids Achelous spinimanus, from the lower Miocene Pirabas Formation, State of Pará, each based on one poorly preserved specimen. The sternum and only part of the chelipeds of the purported ‘A. spinimanus’ are preserved, making the diagnostic characteristics of the species difficult to observe. The material of N. tajinensis is even less well preserved and also in ventral view, which makes the identification unreliable. Thus, we consider both records as doubtful. Likewise, the affiliation of the sole propodus of the cheliped referred to S. costata is dubious.
Note 6. In a recent work, Távora and Dias (2016) reported the occurrence of the swimming crab Portunus oblongus based on relatively poorly preserved cheliped and dorsal carapace material. Although it is possible that their specimens belong to Portunus, their specific assignment to P. oblongus is not justified or discussed, especially given that at least three other species of Portunus 3(4 i.e., P. atecuicitlis, P. haitensis, and P. pirabaensis 3(4, and two of its close relative Callinectes 3(4i.e., C. paraensis and C. reticulatus 3(4, are known from the same strata and age in the state of Pará. Therefore, herein we consider this record as Portunus spp. A second record assigned herein to Portunus spp. corresponds to what they incorrectly called Palaeopinnixa porornata ENT#091;sicENT#093; Távora and Dias (2016) (correct spelling is perornata) following the typographic error in the original publication of the species by Collins and Morris (1976, p. 127). Palaeopinnixa perornata is a crab of the family Hexapodidae 3(4 not Pinnotheridae, as suggested by the Távora and Dias (2016) 3(4, characterized by the unusual reduction of their fifth pair of pereiopods and their corresponding sternite 8, giving the impression of having only three pairs of walking legs (hence Hexapodidae). Hexapodid crabs share a small and wide pea-like carapace with some Pinnotheridae due to convergence (usually no more than a couple centimeters total), which has previously influenced some authors to place fossil hexapodids among pinnotherids (Vía Boada, 1966; Collins and Rasmussen, 1992; Schweitzer and Feldmann, 2001). The large specimen illustrated by Távora and Dias (2016) as Palaeopinnixa perornata corresponds to the ventral carapace of a medium size Portuninae, most likely one of the Portunus or Callinectes species mentioned above. Another taxon reported by Távora and Dias (2016) was incorrectly assigned to Piloslambrus guerini (Parthenopidae). Despite the poor preservation and illustration of that specimen, the dorsal regions, carapace outline, and tuberculation pattern indicate that this fossil is neither conspecific nor congeneric with the extant P. guerini. Herein, we consider it provisionally as Eubrachyura incertae sedis. Some portunid remains illustrated in Brito (1971), as Callinectes ferreirai and C. pirabensis are here considered as Portunus spp.
Note 7. The original description of ‘Archaeopus’ rathbunae does not include images of the holotype, but only a single line drawing (Beurlen, 1965, fig. 4). Based exclusively on the line drawing provided by Beurlen, the specimen does not seem assignable to Archaeopus, casting doubt on its generic placement.
Note 8. A small lot of cheliped fragments in the USNM Paleobiology collections, collected in 1973 from “Pleistocene dune deposits at Ponto Santo Antonio, Fernando de Noronha Island, S. Atlantic Ocean (Brazil)”, is referred to the extant gecarcinid crab Johngarthia lagostoma (Fig. 6). The recent discovery of claw remains of J. lagostoma in Holocene deposits from the Rocas Atoll, nearby the Fernando de Noronha Island (Soares et al., 2016), confirm the presence of the species in the South Atlantic Islands during the Quaternary. These occurrences represent the first fossil records of the genus (Fig. 6).
Note 9. Brito (1972) named Uca antiqua as a subspecies of U. maracoani (i.e., U. maracoani antiqua). Later, Martins-Neto (2001) recognized that both subspecies were different enough to merit independent species status. However, instead of elevating antiqua to the species level, he erected a new species, U. inaciobritoi, to replace it. Following Article 23.3.1. Principle of Priority of the ICZN, Uca antiquaBrito, 1972, is the valid name for the taxon, and U. inaciobritoiMartins-Neto, 2001, is thus a junior synonym, and therefore invalid. Távora (2001) synonymized U. antiqua with U. maracoani based on the study of 96 Uca specimens from Pirabas Formation. However, Távora (2001) did not provide characters to support his point of view. Herein, we maintain both U. antiqua and U. maracoani as valid separate species.
CHILE
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
?Paguroidea incertae sedis ENT#091;Maastrichtian, AlgarroboENT#093; (Schweitzer et al., 2006a).
Infraorder BRACHYURA Latreille, 1802
Superfamily HOMOLOIDEA De Haan, 1833
Homolidae De Haan, 1839
Homolopsis † Bell, 1863
Homolopsis chilensis † Förster and Stinnesbeck, 1987 ENT#091;Maastrichtian, near ConcepciónENT#093;.
Superfamily RANINOIDEA De Haan, 1839
Lyreididae Guinot, 1993
Lyreidinae Guinot, 1993
Lyreidus De Haan, 1841
Lyreidus lebuensis † Feldmann, 1992 ENT#091;Eocene, near LebuENT#093; (as Lyreidus sp. inFeldmann and Chirino-Galvez, 1991).
Raninidae De Haan, 1839
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides araucana † (Philippi, 1887, as Symnista araucana) ENT#091;Eocene, near LebuENT#093;.
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus spinimarginatus † Feldmann, Schweitzer and Encinas, 2005 ENT#091;Miocene, Cardenal CaroENT#093;.
Calappidae De Haan, 1833
Calappilia † A. Milne-Edwards, 1873
?Calappilia chilensis † Feldmann, Schweitzer and Encinas, 2005 ENT#091;Miocene, Cardenal CaroENT#093;.
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Ixa Leach, 1817
Ixa sp. cf. I. cylindrus (Fabricius, 1777) ENT#091;Cenozoic indet., locality unknownENT#093; (Fig. 7) ENT#091;Note 1ENT#093;.
Fossil Eubrachyura from the Cenozoic of Chile, South America. A, B, Leucosoidea: Leucosidae: Ixa sp. cf. I. cylindrus (Fabricius, 1777), MNHN-F.R03449, Cenozoic indet., locality unknown, in dorsal (A) and frontal (B) views. Photos courtesy of Peter Massicard and Sylvain Charbonnier (MNHN). All specimens dry, uncoated.
Superfamily CANCROIDEA Latreille, 1802
Atelecyclidae Ortmann, 1893
Trichopeltarion A. Milne-Edwards, 1880b
Trichopeltarion frassinetti † Feldmann, Schweitzer and Encinas, 2010 ENT#091;late Pliocene, Guafo IslandENT#093;.
Trichopeltarion levis † Casadío et al., 2004 ENT#091;Miocene, Cardenal CaroENT#093; (Feldmann et al., 2005; 2010).
Superfamily CHEIRAGONOIDEA Ortmann, 1893
Cheriagonidae Ortmann, 1893
Pirulella † Feldmann, Schweitzer and Encinas, 2010
Pirulella antipodea † Feldmann, Schweitzer and Encinas, 2010 ENT#091;early Pliocene, Chiloé IslandENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Geryonidae Colosi, 1923
Archaeogeryon † Colosi, 1923 (= Proterocarcinus Feldmann, Schweitzer and Encinas, 2005 )
Archaeogeryon navidad † (Feldmann, Schweitzer and Encinas, 2005) ENT#091;Miocene, Cardenal CaroENT#093; (Feldmann et al., 2010).
Chaceon Manning and Holthuis, 1989
Chaceon quadrata † Feldmann, Schweitzer and Encinas, 2010 ENT#091;Miocene, CorrálENT#093;.
Geryon Krøyer, 1837
Geryon manningiFeldmann, Schweitzer and Encinas, 2010 ENT#091;early Pliocene, Chiloé IslandENT#093;.
Macropipidae Stephenson and Campbell, 1960
Minohellenus † Karasawa, 1990
Minohellenus araucanus † (Philippi, 1887) ENT#091;Miocene, locality indet.ENT#093;.
Portunidae Rafinesque, 1815
Pheophthalmus † Feldmann, Schweitzer and Encinas, 2010
Pheophthalmus mochaensis † Feldmann, Schweitzer and Encinas, 2010 ENT#091;Miocene, Mocha IslandENT#093;.
Superfamily GONEPLACOIDEA MacLeay, 1838
Chasmocarcinidae Serène, 1964
Chasmocarcininae Serène, 1964
Chasmocarcinus Rathbun, 1898
Chasmocarcinus chiloeensis † Feldmann, Schweitzer and Encinas, 2010 ENT#091;Miocene, Chiloé IslandENT#093;.
Superfamily PILUMNOIDEA Samouelle, 1819
Pilumnidae Samouelle, 1819
Pilumninae Samouelle, 1819
Pilumnus Leach, 1816
Pilumnus cucaoensis † Feldmann, Schweitzer and Encinas, 2005 ENT#091;Miocene, Chiloé Island and Cardenal CaroENT#093; (Feldmann et al., 2010).
Subsection THORACOTREMATA Guinot, 1977
Superfamily PINNOTHEROIDEA De Haan, 1833
Pinnotheridae De Haan, 1833
Pinnothereliinae Alcock, 1900a
Pinnixa White, 1846
Pinnixa navidadensis † Feldmann, Schweitzer and Encinas, 2005 ENT#091;Miocene, Cardenal CaroENT#093; (Feldmann et al., 2010).
Note 1. This is the first occurrence of fossil Ixa in South America, and apparently the second record worldwide (Schweitzer et al., 2010). Unfortunately, no detailed geographic or stratigraphic data are associated with the specimen here reported (Fig. 7), or the record listed in Schweitzer et al. (2010).
COLOMBIA
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
?Paguristes Dana, 1851c
?Paguristes sp. ENT#091;late Valanginian, SantanderENT#093; (Fig. 8A) ENT#091;Note 1ENT#093;.
Fossil Anomura and Brachyura from the Cretaceous and Cenozoic of Colombia, South America. A, Anomura: Paguroidea: Diogenidae: ?Paguristes sp. left cheliped, specimen IGM p880851, late Valanginian, Santander. B-J, Brachyura; B, C, Raninoida; B, Palaeocorystoidea: Palaeocorystidae: Joeranina kerri (Luque et al., 2012), dorsal view of holotype IGM p881128, late Aptian of Santander; C, Raninoidea: Raninidae: Raninoidinae: Quasilaeviranina sp., dorsal view of uncatalogued specimen, early-middle Santonian of Boyacá; D-J, Eubrachyura; D, Dorippoidea: Telamonocarcinidae: Telamonocarcinus antiquus Luque, 2015a, dorsal view of holotype IGM p881012, early Albian of Santander. E, Carpilioidea: Palaeoxanthopsidae: Palaeoxanthopsis sp., dorsal view of specimen IGM IGM p881293, Maastrichtian of Santander. F, G, Portunoidea: Portunidae: Portuninae: Portunus oblongus Rathbun, 1920b, from the Miocene of La Guajira; F, dorsal view, specimen MUN-STRI 37322; G, male ventral view, specimen MUN-STRI 37324. H, I, Retroplumoidea: Retroplumidae: Costacopluma sp. from the Maastrichtian of Santander; H, dorsal carapace, specimen IGM p881282; I, dorsal carapace, specimen IGM p881282. J, Eubrachyura incertae sedis, dorsal view of specimen IGM p881262, Paleocene of La Guajira. Photos by Javier Luque. Specimens C, D, J dry, coated with ammonium chloride; all other specimens dry, uncoated.
Infraorder BRACHYURA Latreille, 1802
Superfamily DROMIOIDEA De Haan, 1833
Diaulacidae † Wright and Collins, 1972
Diaulax Bell, 1863
Diaulax rosablanca † Gómez, Bermúdez and Vega, 2015 ENT#091;late Valanginian, SantanderENT#093;.
Diaulax sp. † ENT#091;late Valanginian to Albian, SantanderENT#093;.
Superfamily HOMOLOIDEA De Haan, 1833
Mithracitidae † Števčić, 2005
Mithracites † Gould, 1859
Mithracites takedai † Van Bakel, Guinot, Jagt and Fraaije, 2012b ENT#091;late Aptian, SantanderENT#093;
Superfamily NECROCARCINOIDEA † Förster, 1968
Cenomanocarcinidae † Guinot, Vega and Van Bakel, 2008
Cenomanocarcinus † Van Straelen, 1936
Cenomanocarcinus vanstraeleni † Stenzel, 1945 (as Ophthalmoplax spinosus inFeldmann, Villamil and Kauffman, 1999; and as ?Pinnotheres sp. inFeldmann et al., 1999) ENT#091;late Albian to Campanian of Cundinamarca, Boyacá and TolimaENT#093; (Vega et al., 2007b; 2010) ENT#091;Note 2ENT#093;.
Orithopsidae † Schweitzer, Feldmann, Fam, Hessin, Hetrick, Nyborg and Ross, 2003
Bellcarcinus † Luque, 2014
Bellcarcinus aptiensis † Luque, 2014 (type) ENT#091;late Aptian, Santander and BoyacáENT#093; ENT#091;Note 3ENT#093;.
Colombicarcinus † Karasawa, Schweitzer, Feldmann and Luque, 2014.
Colombicarcinus laevis † Karasawa, Schweitzer, Feldmann and Luque, 2014 (type) ENT#091;late Aptian, Santander and BoyacáENT#093; ENT#091;Note 3ENT#093;.
Planocarcinus † Luque, Feldmann, Schweitzer, Jaramillo and Cameron, 2012
Planocarcinus olssoni † (Rathbun, 1937, as Dakoticancer olssoni) (type) (and as Necrocarcinus inFeldmann et al., 1999; and Orithopsis inVega et al., 2010) ENT#091;late Aptian, Santander and BoyacáENT#093; (Luque et al., 2012) ENT#091;Note 3ENT#093;.
Planocarcinus johnjagti † Bermúdez, Crúz and Vega in Bermúdez et al., 2013 ENT#091;late Aptian, BoyacáENT#093;.
Superfamily PALAEOCORYSTOIDEA † Lőrenthey in Lőrenthey and Beurlen, 1929
Palaeocorystidae † Lőrenthey in Lőrenthey and Beurlen, 1929
Joeranina † Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a
Joeranina kerri † (Luque, Feldmann, Schweitzer, Jaramillo and Cameron, 2012, as Notopocorystes kerri) ENT#091;late Aptian-middle Albian, Santander and BoyacáENT#093; (Fig. 8B).
Joeranina colombiana † Bermúdez, Crúz and Vega in Bermúdez et al., 2013 ENT#091;early-middle Albian, BoyacáENT#093;.
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Quasilaeviranina † Tucker, 1998
Quasilaeviranina sp. † ENT#091;early-middle Santonian to Maastrichtian of Boyacá, Tolima, and CundinamarcaENT#093; (Fig. 8C) ENT#091;Note 4ENT#093;.
Section EUBRACHYURA Saint Laurent, 1980
Eubrachyura indet. (as ?Goneplacoidea incertae sedis, inKiel and Hansen, 2015) ENT#091;Oligocene, CórdobaENT#093; ENT#091;Note 5ENT#093;.
Eubrachyura indet. 2 ENT#091;Paleocene, GuajiraENT#093; (Fig. 8J) ENT#091;Note 6ENT#093;.
Subsection HETEROTREMATA Guinot, 1977
Superfamily DORIPPOIDEA MacLeay, 1838
Telamonocarcinidae † Larghi, 2004
Telamonocarcinus † Larghi, 2004
Telamonocarcinus antiquus † Luque, 2015a ENT#091;early Albian, SantanderENT#093; (Fig. 8D) ENT#091;Note 7ENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Macropipidae Stephenson and Campbell, 1960
Ophthalmoplax † Rathbun, 1935b
Ophthalmoplax andina † Guzmán, Bermúdez, Gómez-Cruz and Vega, 2016 ENT#091;Campanian, BoyacáENT#093; ENT#091;Note 6ENT#093;.
Ophthalmoplax brasiliana † (Maury, 1930) ENT#091;junior synonym O. triambonatusFeldmann and Villamil, 2002ENT#093; ENT#091;Maastrichtian, BoyacáENT#093; ENT#091;Note 8ENT#093;.
Portunidae Rafinesque, 1815
Portuninae Rafinesque, 1815
Callinectes Stimpson, 1862
Callinectes reticulatus † Rathbun, 1918 (claw) ENT#091;Oligocene, AntioquiaENT#093;.
Portunus Weber, 1795
Portunus oblongus † Rathbun, 1920b ENT#091;Miocene, La GuajiraENT#093; (Fig. 8F, G) ENT#091;Note 9ENT#093;.
Superfamily RETROPLUMOIDEA Gill, 1894
Retroplumidae Gill, 1894
Costacopluma † Collins and Morris, 1975
Costacopluma spp. † ENT#091;Coniacian to Maastrichtian of Tolima, Cundinamarca, Boyacá and SantanderENT#093; (Fig. 8H, I) ENT#091;Note 10ENT#093;.
Superfamily TRICHODACTYLOIDEA H. Milne Edwards, 1853
Trichodactylidae H. Milne Edwards, 1853
Dilocarcininae Pretzmann, 1978
Sylviocarcinus H. Milne Edwards, 1853
Sylviocarcinus piriformis (Pretzmann, 1968) ENT#091;early Miocene, TolimaENT#093; (Fig. 4B) ENT#091;Note 11ENT#093;.
Superfamily CARPILIOIDEA Ortmann, 1893
Palaeoxanthopsidae † Schweitzer, 2003
Palaeoxanthopsis † Beurlen, 1958b
Palaeoxanthopsis sp. † ENT#091;Maastrichtian, SantanderENT#093; (Fig. 8E) ENT#091;Note 12ENT#093;.
Note 1. This represents the first record of anomuran crabs from Colombia, and one of the oldest decapod crustaceans for northern South America (Fig. 8A). Furthermore, if its generic affinities are confirmed, this would be the earliest record for the genus, extending its temporal and geographic record from the late Cretaceous of North America to the early Cretaceous of northern South America (~135 Ma).
Note 2. The species reported by Feldmann et al. (1999) as Ophthalmoplax spinosus from the Turonian of Colombia does not belong to Ophthalmoplax or any eubrachyuran crab. The material represents a taxon congeneric with Cenomanocarcinus, as suggested by Vega et al. (2007b). The specimen of ?Pinnotheres sp. inFeldmann et al. (1999) seems to correspond also to a poorly preserved carapace of Cenomanocarcinus sp.
Note 3. Colombicarcinus and Planocarcinus share several traits with Necrocarcinidae and Orithopsidae, but the absence of ventral and appendicular characters obscures their familial affinities. A recent revision of these taxa suggests that they may represent either basal orithopsids, or even representatives of distinctive orithopsid sister clades likely related to Bellcarcinus and similar forms (Schweitzer et al., 2016). Currently, the age of Planocarcinus olssoni is considered as late Aptian based on the youngest rocks outcropping near the area of collection (Luque et al., 2012), in contrast to the Barremian age originally suggested by Rathbun (1937). Although it is plausible that P. olssoni was already present in the Barremian, the recent discovery of P. olssoni in the late Aptian-early Albian rocks of Boyacá in association with new records of Bellcarcinus aptiensis and Joeranina kerri (J. Luque, pers. obs.) confirms its presence during the latest early Cretaceous (late Aptian-early Albian).
Note 4. This is the first record of the genus Quasilaeviranina for tropical America, and the oldest record worldwide, i.e., early-middle Santonian (~85 Mya), given that all other quasilaeviraninids known so far are Paleocene-Eocene in age (Van Bakel et al., 2012a; Karasawa et al., 2014; Martínez-Díaz et al., 2017; Luque, in prep.).
Note 5. A fragmented dorsal carapace of an Oligocene brachyuran crab was assigned to ?Goneplacoidea inKiel and Hansen (2015, fig. 5G). Investigation of the original material suggests that 1) it was illustrated upside down, and 2) that it is not a goneplacoid but may represent a xanthoid-like species (J. Luque, pers. obs.).
Note 6. This eubrachyuran from the Cerrejón Formation in La Guajija represents the first record of Paleocene decapods in Colombia. A detailed description of this taxon is forthcoming (Luque, in prep.).
Note 7. Telamonocarcinus antiquus is the oldest representative of the family Telamonocarcinidae and the superfamily Dorippoidea yet known, and together with Componocancridae Feldmann et al., 2008, and Tepexicarcinidae Luque, 2015a, represent the earliest confirmed crown- and stem-group Eubrachyura 3(4 or higher true crabs 3(4 known to date (late Albian). Their geographic occurrences suggest that early eubrachyurans could have radiated in the Americas during the early Cretaceous or earlier (Luque, 2015a).
Note 8. The fossil record of Ophthalmoplax was recently reviewed by Vega et al. (2013), who recognized the morphological similarities between O. brasiliana and several records from Colombia and Venezuela previously reported as O. triambonatusFeldmann and Villamil, 2002, and currently synonymized with O. brasiliana (Jagt et al., 2015; Guzmán et al., 2016). Yet, the systematic relationships among species of Ophthalmoplax and related genera need to be revised.
Note 9. This is the first record of Portunus oblongus for Colombia. It is a widespread species found in Miocene rocks of Venezuela, Cuba, Dominican Republic, and Trinidad (e.g., Rathbun, 1920b; Collins and Morris, 1976; Feldmann and Schweitzer, 2004; Varela and Rojas-Consuegra, 2009; Aguilera et al., 2010).
Note 10. Costacopluma is one of the most widespread brachyurans in the Cretaceous of Colombia, occurring in Coniacian to Maastrichtian rocks of Tolima, Cundinamarca, Boyacá, and Santander, and typically associated with raninids, palaeoxanthopsids, and axiidean shrimp (J. Luque, pers. obs.). These are the first reports of Costacopluma and the family Retroplumidae for Colombia, and a detailed description of the species is forthcoming.
Note 11. The specimens of Sylviocarcinus from the Miocene Villavieja Formation of Colombia are represented solely by cheliped fragments, principally pollices, dactyli, and fragments of the palm. Rodriguez (1997) considered these remains conspecific with the extant S. piriformis, which today lives in the same watershed. However, it is unclear if the fossil material indeed belongs to the same species, as no carapaces have been discovered, which makes the verification of its systematic placement difficult.
Note 12. This is the first record of the genus Palaeoxanthopsis and the family Palaexanthopsidae for Colombia. Its Maastrichtian age confirms the wide distribution of the genus and family in the Americas during the late Cretaceous (Rathbun, 1902; Vega et al., 2001b; Schweitzer et al., 2008) (see also under ‘Jamaica’ herein).
ECUADOR
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes proavitus † (Rathbun, 1918) ENT#091;early Miocene, Cuenca BasinENT#093; (Morris, 1973) ENT#091;Note 1ENT#093;.
Portuninae Rafinesque, 1815
Portunus Weber, 1795
Portunus oblongus † Rathbun, 1920b ENT#091;late Miocene, locality indet.ENT#093; (in Collins and Morris, 1976)
Note 1.Necronectes proavitus is known from Miocene deposits of Ecuador, Venezuela, Panama, Puerto Rico, and apparently also Trinidad (Roberts, 1975; Feldmann et al., 1993; Schweitzer et al., 2002; 2006c; Collins et al., 2009c; Cáceres et al., 2016), which indicates a trans-isthmian distribution for the taxon. To our knowledge, this is the only record of fossil crabs from Ecuador.
PERU
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Epialtidae MacLeay, 1838
Epialtinae MacLeay, 1838
Eoinachoides † Van Straelen, 1933b
Eoinachoides latispinosa † Carriol, Muizon and Secretán, 1987 ENT#091;late Miocene, PiscoENT#093; (Fig. 9C).
Fossil Eubrachyura from the Cenozoic of Peru, South America. A, B, Cancroidea: Cancridae: Cancer borealis Stimpson, 1859, early Pliocene of Pisco, dorsal view of specimens MNHN.F.R07746 (A) and MNHN.F.R07748 (B). C, Majoidea: Majidae: Epialtinae: Eoinachoides latispinosa Carriol, Carriol, Muizon and Secretán, 1987, late Miocene of Pisco, holotype, MNHN.F.R70743, dorsal view. D, E, Portunoidea: Geryonidae: Chaceon peruvianus (d’Orbigny, 1842), Miocene of Sasaco (locality unknown), holotype, MNHN.F.B33420, dorsal (D) and ventral (E) views. F-H, Portunidae: Podophthalminae: ?Euphylax sp., Eocene of Pisco, specimen USNM 618318, dorsal view (F); specimen USNM 618319a, dorsal view (G); specimen USNM 618319b, ventral view (H). I-K, Portuninae: Callinectes sp. aff. C. reticulatus Rathbun, 1918, middle ?Oligocene of Piura, specimen USNM 496112a, dorsal (I) and frontal (J) views; specimen USNM 496112b, dorsal view (K). Photos A-D courtesy of Jocelyn Falconnet, Peter Massicard, and Sylvain Charbonnier (MNHN); photos F-K by Javier Luque. Specimens A-D dry, uncoated; F-K dry, coated with ammonium chloride.
Pisinae Dana, 1851b
Libinia Leach, 1815
Libinia peruviensis † Carriol, Muizon and Secretán, 1987 ENT#091;early Pliocene, PiscoENT#093;.
Superfamily CANCROIDEA Latreille, 1802
Cancridae Latreille, 1802
Cancrinae Latreille, 1802
Cancer Linnaeus, 1758
Cancer borealisStimpson, 1862 ENT#091;Miocene-early Pliocene, Pisco and ArequipaENT#093; (Fig. 9A, B).
Superfamily PORTUNOIDEA Rafinesque, 1815
Geryonidae Colosi, 1923
Chaceon Manning and Holthuis, 1989
Chaceon peruvianus † (Orbigny, 1842, as Portunus) ENT#091;Miocene, locality unknownENT#093; (Fig. 9D, E) ENT#091;Note 1ENT#093;.
Portunidae Rafinesque, 1815
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphylax sp. cf. E. callinectias † Rathbun, 1918 ENT#091;Eocene, PiscoENT#093; (Fig. 9F-H) ENT#091;Notes 2, 3ENT#093;.
Portuninae Rafinesque, 1815
Achelous De Haan, 1833
Achelous gibbesii (Stimpson, 1862) ENT#091;Miocene to Pleistocene, Falcón and SucreENT#093; (Aguilera et al., 2010
Callinectes Stimpson, 1862
Callinectes ?reticulatus † Rathbun, 1918 ENT#091;middle Oligocene, PiuraENT#093; (Fig 9I-K) ENT#091;Note 3ENT#093;.
Superfamily TRICHODACTYLOIDEA H. Milne Edwards, 1853
Trichodactylidae H. Milne Edwards, 1853
Trichodactylidae spp. indet. ENT#091;middle Eocene to late Oligocene, Loreto and San MartínENT#093; (Klaus et al., 2017).
Superfamily CARPILIOIDEA Ortmann, 1893
Zanthopsidae † Vía Boada, 1959
Zanthopsis † M'Coy, 1849
Zanthopsis errans † Woods, 1922 ENT#091;Eocene, PiuraENT#093;.
Superfamily HEXAPODOIDEA Miers, 1886
Hexapodidae Miers, 1886
Palaeopinnixa † Vía Boada, 1966
Palaeopinnixa eocenica † (Woods, 1922, as Thaumastoplax) ENT#091;Eocene, PiuraENT#093;.
Note 1. The label associated with the holotype of Chaceon peruvianus (Orbigny, 1842), MNHN.F.B33420 (Fig. 9D, E), indicates that the specimen comes from Miocene rocks outcropping south of the Sasaco basin, likely near Nazca and Arequipa, but any details of its provenance are unknown (Orbigny, 1842, part 4, t. 3, p. 107).
Note 2. Re-examination of three specimens catalogued as ?Callinectes sp., i.e., USNM 618318 (one sample, Fig. 9F), and USNM 618319 (two samples, Fig. 9G, H) from the lowermost upper Eocene Basal Talara shales (Discocyclina peruviana zone) about a mile northwest of Lagunitas, Pisco, Peru, indicates that they belong to Euphylax. To our knowledge, this would represent the first record of the subfamily Podophthalminae and the genus Euphylax for Peru.
Note 3. This is the first record of the subfamily Portuninae and the genus Callinectes for Peru (Fig. 9I-K). The occurrence in Peru of Euphylax and Callinectes in Eocene and Oligocene rocks, and Cancer and Eoinachoides in Miocene and Pliocene deposits (see Carriol et al., 1987), indicate that these genera had a wide trans-isthmian distribution during the Paleogene and Neogene.
VENEZUELA
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Paguristes Dana, 1851c
Paguristes sp. ENT#091;early Miocene, FalcónENT#093; (Aguilera et al., 2010).
Petrochirus Stimpson, 1858
Petrochirus sp. ENT#091;Oligo-Miocene to Pleistocene of Falcón, Lara and SucreENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010).
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides rathbunae † Van Straelen, 1933b ENT#091;late Eocene (Priabonian) to Miocene, Falcón and LaraENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010) ENT#091;Note 1ENT#093;.
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Epialtidae MacLeay, 1838
Epialtinae MacLeay, 1838
Eoinachoides † Van Straelen, 1933b
Eoinachoides senni † Van Straelen, 1933b (type) ENT#091;middle Eocene to Miocene, FalcónENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010).
Eoinachoides sp. † ENT#091;late Miocene, FalcónENT#093; (Aguilera et al., 2010).
Mithracidae MacLeay, 1838
Mithrax Desmarest, 1823
Mithrax sp. ENT#091;Plio-Pleistocene, Falcón and SucreENT#093; (Aguilera et al., 2010).
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Eriosachila † Blow and Manning, 1996
Eriosachila rathbunae † (Maury, 1930) ENT#091;Eocene to Miocene of Falcón, Lara and ZuliaENT#093; (Van Straelen, 1933b; Feldmann and Schweitzer, 2004; Aguilera et al., 2010).
Eriosachila sp. ENT#091;early Miocene, FalcónENT#093; (Aguilera et al., 2010).
Hepatella Smith, 1869a
Hepatella amazonica † Beurlen, 1958a ENT#091;Miocene, Falcón and LaraENT#093; (Aguilera et al., 2010).
Hepatus Latreille, 1802
Hepatus gronovii Holthuis, 1959 ENT#091;Pleistocene, SucreENT#093; (Aguilera et al., 2010)
Hepatus sp. ENT#091;early Miocene, FalcónENT#093; (Aguilera et al., 2010) ENT#091;Note 2ENT#093;.
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa laraensis † Van Straelen, 1933b ENT#091;middle Eocene to early Miocene, LaraENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010).
Calappa nitida Holthuis, 1958 ENT#091;Pleistocene, SucreENT#093; (Aguilera et al., 2010).
Calappa sp. ENT#091;Oligocene to early Pliocene of Falcón, Lara and SucreENT#093; (Aguilera et al., 2010) ENT#091;Note 2ENT#093;.
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Persephona Leach, 1817
Persephona sp. cf. P. punctata (Linnaeus, 1758) ENT#091;Oligo-Miocene to Pleistocene of Falcón, Lara and SucreENT#093; (Aguilera et al., 2010).
Iliacantha Stimpson, 1871
Iliacantha sp. ENT#091;early to late Miocene, FalcónENT#093; (Aguilera et al., 2010).
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Parthenope Weber, 1795
Parthenope venezuelensis † Van Straelen, 1933b ENT#091;early to late Miocene (not Oligocene), FalcónENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010).
Parthenope sp. ENT#091;early Miocene to Pliocene, Falcón and SucreENT#093; (Aguilera et al., 2010)
Platylambrus Stimpson, 1871
Platylambrus sp. ENT#091;early Miocene to Pliocene, FalcónENT#093; (Aguilera et al., 2010).
Superfamily PORTUNOIDEA Rafinesque, 1815
Macropipidae Stephenson and Campbell, 1960
Ophthalmoplax † Rathbun, 1935b
Ophthalmoplax brasiliana † (Maury, 1930 ENT#091;junior synonym O. triambonatusFeldmann and Villamil, 2002ENT#093;) ENT#091;Maastrichtian, TáchiraENT#093; (Aguilera et al., 2010).
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes proavitus † (Rathbun, 1918) ENT#091;late Miocene, Falcón) (Cáceres et al., 2016)
?Necronectes sp. † ENT#091;Miocene, LaraENT#093; (Aguilera et al., 2010) ENT#091;Note 2ENT#093;.
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphylax sp. ENT#091;Miocene to Pliocene, Falcón and ZuliaENT#093;.
Paraeuphylax † Varela and Schweitzer, 2011
Paraeuphyplax cubaensis † Varela and Schweitzer, 2011 ENT#091;early Miocene, ZuliaENT#093; (Fig. 10I) ENT#091;Note 3ENT#093;.
Fossil Eubrachyura from the Cenozoic of Peru and Venezuela. A-D, eubrachyuran cheliped remains indet., MNHN-F.A57960, from the late Miocene of El Jahuay, Peru. A, B, left propodus, outer (A) and inner (B) views; C, D, malformed dactylus, upper (C) and lower occlusal (D) views. E-H, Portunoidea: Portunidae: Portuninae: Callinectes Stimpson, 1862, from the Miocene of Zulia, Venezuela; E, F, Callinectes sp. cf. C. reticulatus Rathbun, 1918, left cheliped, USNM 618310, in outer (E) and inner (F) views; G, H, Callinectes sp. cf. C. declivis Rathbun, 1918, USNM 618306, dorsal carapace in dorsal (G) and ventral (H) views. I, Podophthalminae: Paraeuphylax cubaensis Varela and Schweitzer, 2011, USNM 618317, early Miocene of Zulia, Venezuela, dorsal carapace. J-M, Hexapodoidea: Hexapodidae: Palaeopinnixa Vía Boada, 1966, from the Miocene of Falcón, Venezuela; J, K, Palaeopinnixa sp., USNM 618314, in frontal (J) and dorsal (K) views with a circular hole in left branchial region (see text, Panama, Note 3). L, M, Palaeopinnixa sp., USNM 618316, in frontal (L) and ventral (M) views. Photos A-D courtesy of Jocelyn Falconnet, Peter Massicard, and Sylvain Charbonnier (MNHN); photos E-M by Javier Luque. Specimens A-D, L, M dry, uncoated; E-K dry, coated with ammonium chloride.
?Saratunus † Collins, Lee and Noad, 2003
?Saratunus sp. † ENT#091;early Miocene, FalcónENT#093; (Aguilera et al., 2010) ENT#091;Note 2ENT#093;.
Portuninae Rafinesque, 1815
Callinectes Stimpson, 1862
Callinectes reticulatus † Rathbun, 1918 (claws only) ENT#091;early Miocene, ZuliaENT#093; (Fig. 10E, F).
Callinectes sp. cf. C. declivis † Rathbun, 1918 ENT#091;Miocene, ZuliaENT#093; (Fig. 10G, H).
Portunus Weber, 1795
Portunus gibbesii (Stimpson, 1862) ENT#091;Miocene to Pleistocene, Falcón and SucreENT#093; (Aguilera et al., 2010)
Portunus oblongus † Rathbun, 1920b ENT#091;Miocene, Falcón and LaraENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010).
Portunus spp. ENT#091;early Miocene to Pliocene, FalcónENT#093; (Aguilera et al., 2010).
Scylla De Haan, 1833
Scylla sp. ENT#091;Miocene to Pliocene, Falcón and LaraENT#093; (Aguilera et al., 2010) ENT#091;Note 2ENT#093;.
Superfamily RETROPLUMOIDEA Gill, 1894
Retroplumidae Gill, 1894
Costacopluma † Collins and Morris, 1975
Costacopluma bifida † Collins, Higgs and Cortitula, 1994 ENT#091;Paleocene, ZuliaENT#093;
Costacopluma sp. † ENT#091;Paleocene, TrujilloENT#093; (Aguilera et al., 2010) ENT#091;Note 2ENT#093;.
Superfamily GONEPLACOIDEA MacLeay, 1838
Chasmocarcinidae Serène, 1964
Chasmocarcininae Serène, 1964
Falconoplax † Van Straelen, 1933b
Falconoplax kugleri † Van Straelen, 1933b (type) ENT#091;middle-late Eocene to early Miocene, Lara and FalcónENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010).
Superfamily HEXAPODOIDEA Miers, 1886
Hexapodidae Miers, 1886
Palaeopinnixa † Vía Boada, 1966
Palaeopinnixa perornata † Collins and Morris, 1976 ENT#091;early Miocene (not Oligocene), Falcón and LaraENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010) (Fig. 10J-M) ENT#091;Note 4ENT#093;.
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Eurytium Stimpson, 1859
Eurytium sp. ENT#091;Pliocene, FalcónENT#093; (Aguilera et al., 2010)
Note 1. Feldmann and Schweitzer (2004) commented that the association of Raninoides rathbunae with taxa like Eriosachila and Falconoplax suggests a Miocene age for the assemblage, thus casting doubts on its Eocene age. We concur. However, we tentatively maintain the stratigraphic record of R. rathbunae as ?Eocene - Miocene until new material with clearer stratigraphic information becomes available.
Note 2. Unidentified cheliped remains by Aguilera et al. (2010) have been tentatively assigned to Scylla sp. ENT#091;MioceneENT#093;, ?Necronectes ENT#091;MioceneENT#093; and doubtfully Saratunus ENT#091;MioceneENT#093;. The presence of Necronectes in Venezuela has been recently confirmed (Cáceres et al., 2016). Scylla is known from Puerto Rico and Brazil, so its occurrence in Venezuela and other countries in tropical America would not be unexpected. Aguilera et al. (2010) also reported on the occurrence of indeterminate species of Calappa ENT#091;Miocene to PleistoceneENT#093;, Hepatus ENT#091;MioceneENT#093;, and Costacopluma ENT#091;PaleoceneENT#093;; all three are known from other Venezuelan localities of similar age.
Note 3. A specimen in the USNM Paleobiology collections from the early Miocene of the Zulia State, is herein assigned to Paraeuphylax cubaensis (Fig. 10I). Paraeuphylax superficially resembles Saratunus and Euphylax; two genera previously reported from the Miocene of Falcón and the Pliocene San Gregorio Formation, Venezuela (Aguilera et al., 2010). However, Paraeuphylax differs from these genera by its wider carapace, wider orbits (nearly one-third the carapace width), narrower rostrum (one-tenth the carapace width), and the presence of eight anterolateral spines excluding the outer orbital one (Collins et al., 2003; Varela and Schweitzer, 2011).
Note 4. A specimen of Palaeopinnixa perornata herein illustrated (Fig. 10J, K) has a circular hole over the left branchial region of the carapace. It superficially resembles a predatory drill hole (see Klompmaker et al., 2013b), but it is more circular than bacterial lesions (Klompmaker et al., 2016a). Additional study is needed to check whether the hole penetrates (part of) the cuticle or only the internal mold.
Central America and southern North America
BELIZE
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily PORTUNOIDEA Rafinesque, 1815
Carcineretidae † Beurlen, 1930
Carcineretes † Withers, 1922
Carcineretes planetarius † Vega, Feldmann, Ocampo and Pope, 1997 ENT#091;Maastrichtian, Albion IslandENT#093;.
COSTA RICA
Infraorder ANOMURA MacLeay, 1838
Superfamily GALATHEOIDEA Samouelle, 1819
Porcellanidae Haworth, 1825
Pachycheles Stimpson, 1858
Pachycheles latus † Rathbun, 1918 ENT#091;Pliocene, Port LimónENT#093; (Fig.11A).
Fossil Anomura and Brachyura from Costa Rica and Panama, Central America. A, B, Anomura: Galatheoidea: Porcellanidae: A, Pachycheles latus Rathbun, 1918, holotype, USNM 324264, Pliocene of Port Limón, Costa Rica; B, Petrolisthes avitus Rathbun, 1918, holotype USNM 324266, Pliocene of Port Limón, Costa Rica. C, D, Brachyura: Raninoidea: Raninidae: Raninoidinae: Raninoides sp. cf. R. benedicti Rathbun, 1935a, late Miocene of Panama; C, UF 210170, dorsal carapace, Chiriquí; D, UF 274886, dorsal carapace, Colón. E, Dorippoidea: Goniochelidae: Goniochele armata Rathbun, 1918, holotype, USNM 324259, dactylus, early Miocene of the Panama Canal. F-I, Goneplacoidea: Euryplacidae; F, Euryplax culebrensis Rathbun, 1918, holotype, USNM 324226, right cheliped propodus, early Miocene of the Panama Canal; G-I, Euryplax sp., UF 262570, early Miocene of the Panama Canal, in frontal (G), dorsal (H), and ventral (I) views. J-Q, Chasmocarcinidae: Falconoplax kugleri Van Straelen, 1933b, early Miocene of the Panama Canal; J, K, UF 260866, in dorsal (J) and ventral (K) views; L-N, UF 262570, in frontal (L), dorsal (M), and ventral (N) views; O-Q, UF 260866, in frontal (L), dorsal (M), and ventral (N) views. Photos A, B, E, F courtesy of Rodney Feldmann (KSU); photos C, D, G-Q courtesy of Sean Roberts (FLMNH). Specimens A, B, E, F dry, coated with ammonium chloride; C, D, G-Q dry, uncoated.
Petrolisthes Stimpson, 1858
Petrolisthes avitus † Rathbun, 1918 ENT#091;Pliocene, Port LimónENT#093; (Fig.11B).
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides sp. cf. R. benedictiRathbun, 1935a ENT#091;?early Pleistocene, Burica PeninsulaENT#093;.
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Mithracidae MacLeay, 1838
Thoe Bell, 1836
Thoe asperoides † Collins and Todd in Todd and Collins, 2005 ENT#091;late Pliocene to early Pleistocene, LimónENT#093;.
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus lineatinus † Collins and Todd in Todd and Collins, 2005 ENT#091;late Pliocene to early Pleistocene, Puntarenas).
Hepatus biformis † Collins in Todd and Collins, 2005 ENT#091;early Pliocene, PuntarenasENT#093;.
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa costaricana † Rathbun, 1918 ENT#091;Pliocene; Puerto LimónENT#093; (Todd and Collins, 2005).
Cryptosoma Brullé, 1839
Cryptosoma bairdii (Stimpson, 1862) ENT#091;early Pleistocene, PuntarenasENT#093; (Todd and Collins, 2005).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Leucosilia Bell, 1855
Leucosilia bananensis † Rathbun, 1918 ENT#091;Pliocene, Limón)ENT#093; (Todd and Collins, 2005).
Speleophorus A. Milne-Edwards, 1865
Speleophorus ?subcircularis † Collins and Todd in Todd and Collins, 2005 ENT#091;early Pleistocene, PuntarenasENT#093; (Todd and Collins, 2005).
Persephona Leach, 1817
Persephona sp. cf. P. enigmaticaCollins and Todd in Todd and Collins, 2005 ENT#091;late Pliocene, LimónENT#093;.
Persephona sp. ENT#091;Limón, PuntarenasENT#093; (Todd and Collins, 2005).
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Platylambrus Stimpson, 1871
Platylambrus spinulatus † Collins and Todd in Todd and Collins, 2005 ENT#091;early Pleistocene, LimónENT#093;
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphyplax callinectias † Rathbun, 1918 ENT#091;early Pliocene (not Miocene), LimónENT#093; (Todd and Collins, 2005).
Euphyplax fortis † Rathbun, 1918 ENT#091;early Pliocene (not Miocene), LimónENT#093; (Todd and Collins, 2005).
Euphyplax maculatus † Collins and Todd in Todd and Collins, 2005 (claw fragments) ENT#091;early Pleistocene, PuntarenasENT#093;.
Sandomingia † Rathbun, 1919
Sandomingia yaquiensis † Rathbun, 1919 ENT#091;late Pliocene, LimónENT#093;.
Portuninae Rafinesque, 1815
Callinectes Stimpson, 1862
Callinectes declivis † Rathbun, 1918 (fingers only) ENT#091;early Pliocene to early Pleistocene, LimónENT#093; (Todd and Collins, 2005).
Portunus Weber, 1795
Portunus gabbi † Rathbun, 1919 ENT#091;Pliocene, LimónENT#093;.
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Lophopanopeus Rathbun, 1898
Lophopanopeus maculoides † Collins and Todd in Todd and Collins, 2005 ENT#091;late Pliocene, LimónENT#093;.
Xanthidae MacLeay, 1838
Actaeinae Alcock, 1898
Heteractaea Lockington, 1877
Heteractaea lunata (H. Milne Edwards and Lucas, 1843) ENT#091;late Pliocene to early Pleistocene, LimónENT#093;.
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Cardisoma Latreille, 1828
Cardisoma guanhumiLatreille, 1828 ENT#091;late Pliocene to early Pleistocene, LimónENT#093;.
HONDURAS
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection THORACOTREMATA Guinot, 1977
Superfamily OCYPODOIDEA Rafinesque, 1815
Ocypodidae Rafinesque, 1815
Uca Leach, 1814
Uca sp. aff. U. ornata or U. insignis (as U. ‘marinae’Domínguez Alonso, 2008) ENT#091;Plio-Pleistocene, CholutecaENT#093; ENT#091;Note 1ENT#093;.
Note 1. Following the work of Dominguez (2008), Luque et al. (in press) found that the Honduras material seems to be close to U. ornata, except for the similarity between the ornamentation on the merus of the major claw of some male U. ‘marinae’ and male U. insignis. However, the Honduras material differs from U. insignis in several aspects, which it shares with extant and fossil U. ornata. Alternatively, since U. ornata and U. insignis can be found in sympatry today, it is possible that the fossil assemblage might comprise individuals from both species (Luque et al., in press).
PANAMA
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Dardanus Paul’son, 1875
Dardanus biordines † Collins and Todd in Todd and Collins, 2005 ENT#091;early Pliocene, Bocas del ToroENT#093;.
Petrochirus Stimpson, 1858
Petrochirus bouvieri † Rathbun, 1918 ENT#091;late Miocene to late Pliocene, Bocas del Toro, Colón and DariénENT#093;.
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Ranininae De Haan, 1839
?Ranina Lamarck, 1801
?Ranina sp. ENT#091;Miocene, ChiriquíENT#093;.
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides sp. cf. R. benedictiRathbun, 1935a ENT#091;late Miocene of Bocas del Toro, Chiriquí and ColónENT#093; (Fig. 11C, D).
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily DORIPPOIDEA MacLeay, 1838
?Goniochelidae † Schweitzer and Feldmann, 2011
?Goniochele † Bell, 1858
?Goniochele armata † Rathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093; (Fig. 11E) ENT#091;Note 1ENT#093;.
Superfamily MAJOIDEA Samouelle, 1819
Mithracidae MacLeay, 1838
Thoe Bell, 1836
Thoe asperoides † Collins and Todd in Todd and Collins, 2005 ENT#091;late Miocene, Bocas del ToroENT#093;.
Oregoniidae Garth, 1958
HyasLeach, 1814 ENT#091;inLeach, 1813-1815ENT#093;
Hyas sp. ENT#091;late Miocene, DariénENT#093; (Todd and Collins, 2005).
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Eriosachila † Blow and Manning, 1996
Eriosachila terryi † (Rathbun, 1937, as Zanthopsis terryi) ENT#091;late Eocene, PanamaENT#093;.
Hepatus Latreille, 1802
Hepatus biformis † Collins and Todd in Todd and Collins, 2005 ENT#091;late Miocene, Bocas del ToroENT#093;.
Hepatus chiliensis H. Milne Edwards, 1837 ENT#091;Pleistocene, Panama Canal ZoneENT#093;.
Hepatus lineatinus † Collins and Todd in Todd and Collins, 2005 ENT#091;middle Miocene to late Pliocene, Bocas del Toro and ColónENT#093;.
Hepatus sp. ENT#091;middle Holocene, Bahía de PanamáENT#093; (Portell et al., 2012; Klompmaker et al., 2016b; Luque et al., in press).
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa flammea (Herbst, 1794) ENT#091;late Miocene to Pleistocene, ColónENT#093;.
Calappa zurcheri † Bouvier, 1899 ENT#091;Miocene, unknown provenanceENT#093; (Fig. 12M) ENT#091;Note 2ENT#093;.
Fossil Eubrachyura from Panama, Central America. A-L, Hexapodoidea: Hexapodidae: Palaeopinnixa prima (Rathbun, 1918), early Miocene of the Panama Canal; A-C, UF 217685; D-F, UF 219754; G-I, UF 219750; J-L, UF 219751; specimens in frontal (A, D, G, J), dorsal (B, E, H, K), and ventral (C, F, I, L) views. M, Calappoidea: Calappidae: Calappa zurcheri Bouvier, 1899, holotype, MNHN-F.R03770, dorsal carapace, Miocene of Panama, locality unknown. N, O, Portunoidea: Portunidae: Podophthalminae: ?Euphylax sp., UF 174079, Quaternary, Pacific of Panama, dredged from a depth of ~50.0 meters, in dorsal (N) and ventral (O) views. P, Necronectinae: Necronectes proavitus (Rathbun, 1918), hypotype, USNM 371312, Miocene of Colón, dorsal carapace. Q, Pinnotheroidea: Pinnotheridae: Pinnotherinae: Pinnotheres sp., UF 115397, early Miocene Panama Canal, dorsal carapace. R, Ocypodoidea: Ocypodidae: Ocypodinae: Uca ornata Smith, 1870, USNM 618320, Holocene, Pacific of Panama, dorsal carapace. Photos A-L, N, O courtesy of Sean Roberts (FLMNH); photo M courtesy of Jocelyn Falconnet, Peter Massicard, and Sylvain Charbonnier (MNHN); photos P-R by Javier Luque. Specimens A-O, Q, R dry, uncoated; P dry, coated with ammonium chloride.
Calappa sp. ENT#091;early late Pliocene, Bocas del ToroENT#093; (Todd and Collins, 2005).
Calappella † Rathbun, 1918
Calappella quadrispina † Rathbun, 1918 (type) ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.
Cryptosoma Brullé, 1839
Cryptosoma bairdii (Stimpson, 1862) ENT#091;early late Pliocene, Bocas del ToroENT#093; (Todd and Collins, 2005).
Mursia Desmarest, 1823
Mursia macdonaldi † Rathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.
Mursia obscura † Rathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.
Mursilia † Rathbun, 1918
Mursilia ecristata † Rathbun, 1918 ENT#091;early to middle Miocene, ColónENT#093;.
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Iliacantha Stimpson, 1871
Iliacantha panamica † Collins and Todd in Todd and Collins, 2005 ENT#091;late Miocene, Bocas del ToroENT#093;.
Iliacantha sp. ENT#091;late Pliocene, Bocas del ToroENT#093; (Todd and Collins, 2005).
Leucosilia Bell, 1855
Leucosilia bananensis † Rathbun, 1918 ENT#091;middle-late Miocene, Bocas del Toro and ColónENT#093;.
Leucosilia jurinii (Saussure, 1853) ENT#091;Pleistocene, ColónENT#093; (Rathbun, 1918).
Leucosilia sp. cf. L. jurini ENT#091;Quaternary, Bahía de PanamáENT#093; (Luque et al., in press).
Persephona Leach, 1817
Persephona enigmatica † Collins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.
Persephona manningi † Collins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.
Speloeophorus A. Milne-Edwards, 1865
Speloeophorus subcircularis † Collins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Parthenope Weber, 1795
Parthenope panamensis † Rathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.
Parthenope pleistocenica † Rathbun, 1918 ENT#091;Pleistocene, ColónENT#093;.
Platylambrus Stimpson, 1871
Platylambrus spinulatus † Collins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.
Superfamily CANCROIDEA Latreille, 1802
Cancridae Latreille, 1802
Cancrinae Latreille, 1802
Cancer Latreille, 1802
Cancer santosi † (Rathbun, 1937, as Lobocarcinus santosi) ENT#091;late Eocene, Los SantosENT#093;.
Superfamily CHEIRAGONOIDEA Ortmann, 1893
Cheriagonidae Ortmann, 1893
Montezumella † Rathbun, 1930
Montezumella casayetensis † Rathbun, 1937 ENT#091;late Oligocene or early Miocene, Bahía de PanamáENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes proavitus † (Rathbun, 1918, as Gatunia proavita) ENT#091;early Miocene, ColónENT#093; (Fig. 12N).
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphylax callinectias † Rathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.
Euphylax maculatus † Collins and Todd in Todd and Collins, 2005 ENT#091;middle-late Miocene to late Pliocene, Bocas del Toro, Colón, and DariénENT#093;.
Euphylax sp. ENT#091;Quaternary, Panama Province (dredged)ENT#093; (Fig. 12 N, O).
Sandomingia † Rathbun, 1919
Sandomingia yaquiensis † Rathbun, 1919 ENT#091;early to late Pliocene, Bocas del ToroENT#093;.
Portuninae Rafinesque, 1815
Callinectes Stimpson, 1862
Callinectes declivis † Rathbun, 1918 ENT#091;late Eocene to early Miocene, Panama CanalENT#093;.
Callinectes reticulatus † Rathbun, 1918 ENT#091;early Miocene, Panama CanalENT#093;.
Callinectes sp. cf. C. arcuatusOrdway, 1863 ENT#091;Quaternary, Bahía de PanamáENT#093; (Portell et al., 2012; Luque et al., in press).
Portunus Weber, 1795
Portunus gabbi † Rathbun, 1919 ENT#091;middle-late Miocene, DariénENT#093;.
Portunus spp. cf. P. tenuis † Rathbun, 1919 ENT#091;Pleistocene, ChiriquíENT#093; ENT#091;Note 3ENT#093;.
Superfamily ERIPHIOIDEA MacLeay, 1838
Eriphiidae MacLeay, 1838
Eriphia Latreille, 1817
Eriphia sp. aff. E. squamataStimpson, 1859 ENT#091;Quaternary, Bahía de PanamáENT#093; (Luque et al., in press).
Platyxanthidae Guinot, 1977
Platyxanthus A. Milne-Edwards, 1863
Platyxanthus sp. ENT#091;late Miocene, Bocas del ToroENT#093; (Todd and Collins, 2005).
Superfamily GONEPLACOIDEA MacLeay, 1838
Chasmocarcinidae Serène, 1964
Chasmocarcininae Serène, 1964
Falconoplax † Van Straelen, 1933b
Falconoplax kugleri † Van Straelen, 1933b ENT#091;early Miocene, Panama CanalENT#093; (Fig. 11J-Q) ENT#091;Note 3ENT#093;.
Euryplacidae Stimpson, 1871
Euryplax Stimpson, 1859
? Euryplax culebrensis † Rathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093; (Fig. 11F) ENT#091;Note 5ENT#093;.
Euryplax sp. ENT#091;early Miocene, Panama CanalENT#093; (Fig. 11G-I) ENT#091;Note 5ENT#093;.
Superfamily HEXAPODOIDEA Miers, 1886
Hexapodidae Miers, 1886
Palaeopinnixa † Vía Boada, 1966
Palaeopinnixa prima † (Rathbun, 1918, as Thaumastoplax prima) ENT#091;early Miocene (not Oligocene), Panama CanalENT#093; (Fig. 12A-L) ENT#091;Note 6ENT#093;.
Superfamily PILUMNOIDEA Samouelle, 1819
Pilumnidae Samouelle, 1819
Pilumninae Samouelle, 1819
Pilumnus Leach, 1816
Pilumnus sp. ENT#091;Pliocene - early Pleistocene, Bocas del Toro and ChiriquíENT#093; (Todd and Collins, 2005).
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Eurytium Stimpson, 1859
Eurytium crenulatum † Rathbun, 1918 ENT#091;Pleistocene, ColónENT#093; ENT#091;Note 7ENT#093;.
Panopeus H. Milne Edwards, 1834
Panopeus antepurpureus † Rathbun, 1918 ENT#091;Pleistocene, Colón and DariénENT#093;
Panopeus chilensisMilne Edwards and Lucas, 1843 (claw fragment) ENT#091;?early Pleistocene, Bocas del ToroENT#093; (Todd and Collins, 2005).
Xanthidae MacLeay, 1838
Actaeinae Alcock, 1898
Heteractaea Lockington, 1877
Heteractaea lunata (H. Milne Edwards and Lucas, 1843) ENT#091;late Pliocene, Bocas del ToroENT#093;.
Xanthinae MacLeay, 1838
Micropanope Stimpson, 1871
Micropanope sp. ENT#091;early late Pliocene, Bocas del ToroENT#093; (Todd and Collins, 2005).
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Cardisoma Latreille, 1828
Cardisoma crassumSmith, 1870 ENT#091;Quaternary, Bahía de PanamáENT#093; (Portell et al., 2012; Luque et al., in press) ENT#091;Note 8ENT#093;.
Grapsidae MacLeay, 1838
Grapsus Lamarck, 1801
Grapsus sp. aff. G. grapsus (Linnaeus, 1758) ENT#091;Quaternary, Bahía de PanamáENT#093; (Portell et al., 2012; Luque et al., in press) ENT#091;Note 8ENT#093;.
Superfamily OCYPODOIDEA Rafinesque, 1815
Ocypodidae Rafinesque, 1815
Ocypodinae Rafinesque, 1815
Uca Leach, 1814
Uca ornataSmith, 1870 ENT#091;Quaternary, Bahía de PanamáENT#093; (Portell et al., 2012; Luque et al., in press) ENT#091;Note 9ENT#093;.
Uca sp. ENT#091;as U. macrodactyla inRathbun, 1918) ENT#091;Note 9ENT#093;.
Superfamily PINNOTHEROIDEA De Haan, 1833
Pinnotheridae De Haan, 1833
Pinnothereliinae Alcock, 1900a
Pinnixa White, 1846
Pinnixa sp. ENT#091;early Miocene, Panama Canal (Fig. 12Q) ENT#091;Note 10ENT#093;.
Note 1. Goniochele is the type genus of Goniochelidae, a monotypic eubrachyuran family apparently akin with Dorippoidea (Schweitzer and Feldmann, 2011; Guinot et al., 2013; Luque, 2015a). Its confirmed fossil record is restricted to two species from the Eocene of Europe: Goniochele angulataBell, 1858, and Goniochele madseniCollins and Jakobsen, 2003. A putative third species, Goniochele armataRathbun, 1918 (Fig. 11E), was originally described from the Miocene (not Oligocene) Culebra Formation from the Panama Canal based on the isolated dactylus of a left cheliped. The general elongate triangular shape with armed edges suggested to Rathbun (1918) affinities with Goniochele. Due to the lack of additional material from the Panama Canal, it is hard to confirm Rathbun’s dactylus with Goniochele. Thus, we maintain it in the genus but with uncertainty.
Note 2. Calappa zurcheri is the first described fossil decapod crustacean from Panama. Although Bouvier (1899) reported it as Miocene in age, its exact geographic provenance and stratigraphic context is unknown. Based on our own field observations and the stratigraphic context of recent findings, Bouvier’s specimen could come from the early excavations of the Panama Canal, as we have recovered similar material from the Miocene Culebra and Gatún formations in the area of the recent expansion of the Canal.
Note 3. The material of Portunus tenuis reported by Rathbun (1918) and Todd and Collins (2005) consist of cheliped fragments, making its systematic placement questionable.
Note 4. Falconoplax, despite being one of the most abundant crabs from the Miocene Culebra Formation (hundreds of specimens) (Fig. 11J-Q), is only until recently that has been recognized for Panama (Robins et al. 2016).
Note 5. As noted by Collins et al. (2009c), Rathbun (1918) erected several new genera and species based on fragmentary material, including isolated pollices and dactyli. Euryplax culebrensis does not seem to be an exception. The only goneplacoid-like crabs clearly known from the fossil record of Panama are the abundant Palaeopinnixa prima and Falconoplax kugleri. Both taxa are represented by hundreds of specimens (Robins et al., 2016), often so complete that the appendages are still attached to the body as in life position. It is possible that the holotype of Euryplax culebrensis may represent cheliped material from either P. prima or F. kugleri.
Note 6. Palaeopinnixa prima is similar in shape to the coeval P. perornata from Venezuela, but differs in the carapace outline, the latter having more roundish lateral margins. Palaeopinnixa is, together with Falconoplax, the most abundant crab in the Culebra Formation (Robins et al., 2016).
Note 7. Eurytium crenulatum was described by Rathbun (1918), based on a right dactylus found in Pleistocene sediments near Mount Hope in the Colón Province, Panama. Additional carapace and cheliped material from lower to upper Pliocene rocks of the Limón Province, Costa Rica, has been assigned to E. crenulatum since then (Todd and Collins, 2005; Collins et al., 2009c).
Note 8. The occurrences of the land crab Cardisoma crassum and the Sally Lightfoot crab Grapsus grapsus in the Quaternary of the Pacific Coast of Panama represent the first and second known fossil records of these species to date, respectively (Luque et al., 2015; Luque et al., in press).
Note 9. This middle-Holocene occurrence of Uca ornata accounts for the most complete and abundant fiddler crab fossil record known, constituted by several hundred specimens of juveniles and adults from both sexes (Luque et al., in press). Another fossil fiddler crab from Panama is Uca macrodactylaRathbun, 1918, described based on a single dactylus of an ambulatory leg from Pleistocene deposits near Colón. Crane (1975) commented about the uncertain specific affinities of Rathbun’s fossil, and also the synonymy of U. macrodactyla with U. galapagensis s.l. We concur with Crane (1975) and consider the specimen as an indeterminate species of Uca s.l.
Note 10. This is the first report of fossil pinnotherids from Panama, and the second record of the genus Pinnixa for the Americas, with P. navidadensis from the Miocene of Chile representing the only other record known thus far.
MEXICO
Infraorder ANOMURA MacLeay, 1838
Superfamily AEGLOIDEA Dana, 1852
Aeglidae Dana, 1852
Protaegla † Feldmann, Vega, Applegate and Bishop, 1998b
Protaegla miniscula † Feldmann, Vega, Applegate and Bishop, 1998b (type) ENT#091;Albian, PueblaENT#093; (Vega et al., 2005) (Fig. 13A).
Fossil Anomura and Brachyura from Mexico. A, Anomura: Aegloidea: Aeglidae: Protaegla minuscula Feldmann, Vega, Applegate and Bishop, 1998b, holotype, IGM-6502, late Albian of Puebla. B-L, Brachyura. B, Necrocarcinoidea: Cenomanocarcinidae: Cenomanocarcinus vanstraeleni Stenzel, 1945, hypotype, MUZ-801, Turonian of Múzquiz, Coahuila; C, Raninoidea: Raninidae: Lophoranina cristaspina Vega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a, hypotype, IHNFG-3460, middle Eocene of Copoya, Chiapas; D, Dakoticancroidea: Ibericancridae: Sodakus mexicanus Vega, Feldmann and Villalobos-Hiriart, 1995b, hypotype to be deposited at CPC, early Maastrichtian of Paredón, Coahuila; E, Superfamily uncertain: Tepexicarcinidae: Tepexicarcinus tlayuaensis Feldmann, Vega, Applegate and Bishop, 1998b, hypotype, IGM-6609, late Albian of Puebla; F, Retroplumoidea: Retroplumidae: Costacopluma mexicana Vega and Perrillat, 1989, holotype, IGM-4128, early Maastrichtian of Sierra El Antrisco, Nuevo León, Mexico; G, Cheiragonoidea: Cheiragonidae: Karasawaia markgrafi (Lőrenthey, 1907 1909), hypotype, IHNFG-3030, early Eocene of El Veinte, Chiapas; H-J, Portunoidea; H, Icriocarcinidae: Icriocarcinus xestos Bishop, 1988, hypotype, IGM.6625-2, late Campanian of Punta Santo Tomás, Baja California; I, Carcineretidae: Carcineretes planetarius Vega, Feldmann, Ocampo and Pope, 1997, hypotype, IHNFG-3412, early Maastrichtian of Ocozocoautla, Chiapas; J, Macropipidae: Ophthalmoplax brasiliana (Maury, 1930), hypotype, CPC-881, late Maastrichtian of Arroyo Amargos, Coahuila; K, ?Pinnotheroidea: ?Pinnotheridae: Viapinnixa perrilliatae Vega, Nyborg, Fraaye and Espinosa, 2007a, paratype, IGM-9109, middle Paleocene of La Mesita, Coahuila; L, Grapsoidea: Sesarmidae indet., specimen IHNFG-4991, early Miocene of Simojovel, Chiapas. Photos by Francisco Vega. F and K previously illustrated in Armstrong et al. (2009), and J in Vega et al. (2013). Specimens A-E, G-I, K, L dry, uncoated; F and J dry, coated with ammonium chloride.
Superfamily GALATHEOIDEA Samouelle, 1819
Galatheidae Samouelle, 1819
Galatheidae indet. ENT#091;middle Eocene, Baja California SurENT#093; (Schweitzer et al., 2006b).
Superfamily PAGUROIDEA Latreille, 1802
Paguroidea spp. indet. ENT#091;middle Eocene, Baja California SurENT#093; (Schweitzer et al., 2005).
Diogenidae Ortmann, 1892
?Paguristes Paul’son, 1875
? Paguristes mexicanusVega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a ENT#091;middle Eocene, Chiapas, Baja California SurENT#093; (Schweitzer et al., 2002; 2005; Vega et al., 2008).
Petrochirus Stimpson, 1858
Petrochirus sp. ENT#091;middle Eocene and early Miocene, ChiapasENT#093; (Vega et al., 2008; 2009).
Paguridae Latreille, 1802
Pagurus Fabricius, 1775
Pagurus sp. ENT#091;Albian, PueblaENT#093; (Vega et al., 2005).
Palaeopagurus † Van Straelen, 1925
Palaeopagurus sp. cf. P. pilsbyi † Roberts, 1962 ENT#091;early Maastrichtian, Nuevo LeónENT#093; (Vega et al., 1995b).
Infraorder BRACHYURA Latreille, 1802
Superfamily DROMIOIDEA De Haan, 1833
Dynomenidae Ortmann, 1892
Graptocarcinus † Roemer, 1887
Graptocarcinus muiri † Stenzel, 1944b ENT#091;Albian, San Luis PotosíENT#093;.
Xandarocarcinidae † Karasawa Schweitzer and Feldmann, 2011
Xandarocarcinus †Karasawa, Schweitzer and Feldmann, 2011 (as Xandaros inBishop, 1988)
Xandarocarcinus sternbergi † (Rathbun, 1926) ENT#091;Maastrichtian, Baja Calafornia SurENT#093; (Bishop, 1986; Schweitzer et al., 2002).
Dromioidea incertae sedis
Prehepatus † Rathbun, 1935b
Prehepatus harrisi † Bishop, 1985 ENT#091;early Maastrichtian, Nuevo LeónENT#093; (Vega et al., 1995b).
Prehepatus mexicanus † Schweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.
Superfamily HOMOLOIDEA De Haan, 1833
Homolidae De Haan, 1839
HomolaLeach, 1816 ENT#091;imprint 1815ENT#093;
Homola bajaensis † Schweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.
Zygastrocarcinus † Bishop, 1983a
Zygastrocarcinus carolinasensis † Klompmaker, Flores-Ventura and Vega, 2013a ENT#091;late Campanian, CoahuilaENT#093; (Vega et al., 2016).
Superfamily NECROCARCINOIDEA † Förster, 1968
Cenomanocarcinidae † Guinot, Vega and Van Bakel, 2008
Cenomanocarcinus † Van Straelen, 1936
Cenomanocarcinus vanstraeleni † Stenzel, 1945 ENT#091;Turonian-Coniacian, CoahuilaENT#093; (Vega et al., 2007b; 2010; Garassino et al., 2013) (Fig. 13B).
Cenomanocarcinus sp. † ENT#091;Campanian, GuerreroENT#093; (Vega et al., 2010).
Superfamily PALAEOCORYSTOIDEA † Lőrenthey in Lőrenthey and Beurlen, 1929
Palaeocorystidae † Lőrenthey in Lőrenthey and Beurlen, 1929
Ferroranina † Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a
Ferroranina sp. cf. F. dichrous † (Stenzel, 1945) ENT#091;Turonian, CoahuilaENT#093; (Vega et al., 2007b).
Superfamily RANINOIDEA De Haan, 1839
Lyreididae Guinot, 1993
Macroacaeninae † Karasawa, Schweitzer, Feldmann and Luque, 2014
Macroacaena † Tucker, 1998
Macroacaena venturai † Vega, Nyborg, Fraaye and Espinosa, 2007a ENT#091;middle Paleocene (Selandian), Chiapas and CoahuilaENT#093;.
Marylyreidinae † Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a
Bournelyreidus † Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a
Bournelyreidus oaheensis † (Bishop, 1978, as Raninella oaheensis) ENT#091;early Maastrichtian, CoahuilaENT#093; (Vega et al., 2016).
Raninidae De Haan, 1839
Raninidae spp. indet. ENT#091;Turonian, CoahuilaENT#093; (Vega et al., 2007b).
Cyrtorhininae Guinot, 1993
Claudioranina † Karasawa, Schweitzer, Feldmann and Luque, 2014
Claudioranina latacantha † Martínez, Aguillón, Luque and Vega, 2017 ENT#091;middle Paleocene (Selandian), CoahuilaENT#093;.
Notopodinae Serène and Umali, 1972
Notopus De Haan, 1841
Notopus minutus † Vega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a ENT#091;Ypresian, ChiapasENT#093;.
Ranininae De Haan, 1839
Lophoranina † Fabiani, 1910
Lophoranina bishopi † Squires and Demetrion, 1992 ENT#091;Ypresian and Eocene indet., Baja California, SurENT#093; (Schweitzer et al., 2002; 2007b).
Lophoranina cristaspina † Vega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a ENT#091;middle Eocene and Ypresian, Baja California Sur; Ypresian, ChiapasENT#093; (Hernández-Monzón et al., 2007; Vega et al., 2008) (Fig. 13C).
?Ranina Lamarck, 1801
? Ranina berglundi † Squires and Demetrion, 1992 ENT#091;middle Eocene and Ypresian, Baja California SurENT#093; (Schweitzer et al., 2006b; Vega et al., 2008; Pasini and Garassino, 2017).
?Ranina sp. † ENT#091;Eocene, Baja California SurENT#093; (Rathbun, 1930)
Vegaranina † Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a
Vegaranina precocia † (Feldmann, Vega, Tucker, García-Barrera and Avendaño, 1996, as Lophoranina precocious) (type) ENT#091;Maastrichtian, ChiapasENT#093;.
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Notopoides Henderson, 1888
Notopoides exiguus † Beschin, Busulini, De Angeli and Tessier, 1998 ENT#091;Ypresian, ChiapasENT#093; (Vega et al., 2008).
Notopoides sp. † ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Martínez-Díaz et al., 2017).
Quasilaeviranina † Tucker, 1998
Quasilaeviranina sp. cf. Q. arzignanensis † (Beschin, Busulini, De Angeli and Tessier, 1998) ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Martínez-Díaz et al., 2017)
Quasilaeviranina sp. cf. Q. ovalis † (Rathbun, 1935b) ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Martínez-Díaz et al., 2017).
Raninoides H. Milne Edwards, 1837
Raninoides acanthocolus † Schweitzer, Feldmann, Gonzalez-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.
Raninoides mexicanus † Rathbun, 1930 ENT#091;Miocene, VeracruzENT#093;.
Raninoides proracanthus † Schweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.
Raninoides treldenaesensis † Collins and Jakobsen, 2003 ENT#091;Ypresian, ChiapasENT#093; (Vega et al., 2008).
Section DAKOTICANCROIDA † Rathbun, 1917
Superfamily DAKOTICANCROIDEA Rathbun, 1917
Dakoticancridae † Rathbun, 1917
Dakoticancer † Rathbun, 1917
Dakoticancer australis † Rathbun, 1935b ENT#091;early Maastrichtian, San Luis Potosí and Nuevo LeónENT#093; (Vega and Feldmann, 1991; Vega et al., 1995a; 2016).
Ibericancridae † Artal, Guinot, Van Bakel and Castillo, 2008
?Sodakus † Bishop, 1978
Sodakus mexicanus † Vega, Feldmann and Villalobos-Hiriart, 1995b ENT#091;early Maastrichtian, Nuevo León and CoahuilaENT#093; (Vega et al., 2016) (Fig. 13D).
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily ?DORIPPOIDEA MacLeay, 1838
Tepexicarcinidae † Luque, 2015a
Tepexicarcinus † Feldmann, Vega, Applegate and Bishop, 1998b
Tepexicarcinus tlayuaensis † Feldmann, Vega, Applegate and Bishop, 1998b ENT#091;Albian, PueblaENT#093; (Vega et al., 2005) (Fig. 13E).
Superfamily MAJOIDEA Samouelle, 1819
Oregoniidae Garth, 1958
Oregonia Dana, 1851a
Majidae Samouelle, 1819
Oregonia spinifera Schweitzer, Feldmann, González-Barba, and Vega, 2002 ENT#091;Oligocene, Baja California SurENT#093;.
?Majidae indet. ENT#091;Maastrichtian, San Luis PotosíENT#093; (Vega et al., 1995a).
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Eriosachila † Blow and Manning, 1996
Eriosachila bajaensis † Schweitzer, Feldmann, González-Barba and Vega, 2002 ENT#091;Eocene (Bartonian), Baja California SurENT#093; (Schweitzer et al., 2007a).
Eriosachila sp. † ENT#091;middle Eocene, ChiapasENT#093; (Vega et al., 2001a).
Hepatella Smith, 1869a
Hepatella amazonica † Beurlen, 1958a ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa flammea (Herbst, 1794) ENT#091;Oligocene, Baja California SurENT#093; (Rathbun, 1930).
Calappa zurcheri † Bouvier, 1899 ENT#091;Oligocene, Veracruz; early Miocene, ChiapasENT#093; (Rathbun, 1930; Vega et al., 2009).
Calappilia † A. Milne-Edwards, 1873
Calappilia hondoensis † Rathbun, 1930 ENT#091;middle Eocene: Lutetian, Chiapas; Priabonian, Baja California SurENT#093; (Vega et al., 2001a; 2008; Schweitzer et al., 2006b; 2007a).
Mursia Leach in Desmarest, 1823
?Mursia sp. ENT#091;middle Miocene, VeracruzENT#093; (Vega et al., 1999).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Iliacantha Stimpson, 1871
Iliacantha panamanica † Collins and Todd in Todd and Collins, 2005 ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Daldorfiinae Ng and Rodríguez, 1986
Daldorfia Rathbun, 1904
Daldorfia salina † Schweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.
Superfamily CANCROIDEA Latreille, 1802
Atelecyclidae Ortmann, 1893
Levicyclus † Schweitzer, Feldmann, Gonzáles-Barba and Vega, 2002
Levicyclus tepetate † Schweitzer, Feldmann, González-Barba, and Vega, 2002 ENT#091;Eocene, Baja California SurENT#093;.
Cancridae Latreille, 1802
Cancrinae Latreille, 1802
Anatolikos Schweitzer and Feldmann, 2000
Anatolikos undecimspinosus † Schweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.
Romaleon Gistel, 1848
Romaleon antennarium (Stimpson, 1856) ENT#091;Pliocene, Baja California SurENT#093;.
Superfamily CHEIRAGONOIDEA Ortmann, 1893
Cheriagonidae Ortmann, 1893
Karasawaia † Vega, Nyborg, Coutiño and Hernández-Monzón, 2008
Karasawaia markgrafi † (Lőrenthey, 1907 ENT#091;German version 1907ENT#091;1909ENT#093;, as Plagiolophus markgrafi) (type) ENT#091;Ypresian, ChiapasENT#093; (Vega et al., 2008) (Fig. 13G).
Montezumella † Rathbun, 1930
Montezumella tubulata † Rathbun, 1930 ENT#091;Eocene: Priabonian, Baja California SurENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Carcineretidae † Beurlen, 1930
Carcineretes † Withers, 1922
Carcineretes planetarius † Vega, Feldmann, Ocampo and Pope, 1997 ENT#091;early Maastrichtian, ChiapasENT#093; (Vega et al., 2001b) (Fig. 13I).
Carcinidae MacLeay, 1838
?Xaiva MacLeay, 1838
?Xaiva sp. † ENT#091;middle Eocene, ChiapasENT#093; (Vega et al., 2008).
Icriocarcinidae † Števčić, 2005
Icriocarcinus † Bishop, 1988
Icriocarcinus xestos † Bishop, 1988 ENT#091;Maastrichtian, Baja California SurENT#093; (Schweitzer et al., 2002; Phillips et al., 2014) (Fig. 13H).
Branchiocarcinus † Vega, Feldmann and Sour-Tovar, 1995a
Branchiocarcinus cornatus † Feldmann and Vega, 1995inVega et al., 1995a ENT#091;Maastrichtian, San Luis PotosíENT#093; (Phillips et al., 2014).
Branchiocarcinus flectus † (Rathbun, 1923b, as Eryma flecta) ENT#091;Maastrichtian, San Luis PotosíENT#093; (Phillips et al., 2014; Vega et al., 2016).
Longusorbiidae † Karasawa, Schweitzer and Feldmann, 2008
Longusorbis † Richards, 1975
Longusorbis eutychius † Schweitzer, Feldmann and Karasawa, 2007c ENT#091;middle Eocene (Bartonian), Baja California SurENT#093;.
Longusorbis quadratus † Fraaije, Vega, Van Bakel and Garibay-Romero, 2006 ENT#091;Cenomanian-Coniacian, GuerreroENT#093;.
Macropipidae Stephenson and Campbell, 1960
Maeandricampus † Schweitzer and Feldmann, 2002
Maeandricampus americanus † (Rathbun, 1930, as Xanthosia americana) ENT#091;Eocene, Baja California SurENT#093;.
Ophthalmoplax † Rathbun, 1935b
Ophthalmoplax brasiliana † (Maury, 1930) (as Mascaranada difuntaensis in Vega and Feldmann, 1991) ENT#091;early to late Maastrichtian, Coahuila and Nuevo LeónENT#093; (Vega et al., 2013; 2016) (Fig. 13J).
Portunidae Rafinesque, 1815
Portunidae indet. ENT#091;middle Eocene, ChiapasENT#093; (Vega et al., 2008).
Portunidae indet. ENT#091;middle Eocene, Baja California SurENT#093; (Schweitzer et al., 2006b).
Portunidae indet. ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes nodosus † Schweitzer, Feldmann, González-Barba and Vega, 2002 ENT#091;Oligocene, Baja California SurENT#093;.
Necronectes tajinensis † Vega et al., 1999 ENT#091;middle Miocene, VeracruzENT#093;.
Necronectes sp. † ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).
Portuninae Rafinesque, 1815
Portunus Weber, 1795
Portunus atecuicitlis † Vega et al., 1999 ENT#091;early to middle Miocene, Chiapas and VeracruzENT#093; (Vega et al., 2009).
Scylla De Haan, 1833
Scylla costata † Rathbun, 1919 ENT#091;age and locality unknownENT#093; (Rathbun, 1930).
Podophthalminae Dana, 1851d
Podophthalmus Lamarck, 1801
?Podophthalmus sp. † ENT#091;late Cretaceous, TamaulipasENT#093; (Rathbun, 1930).
Superfamily RETROPLUMOIDEA Gill, 1894
Retroplumidae Gill, 1894
Archaeopus † Rathbun, 1908
Archaeopus mexicanus † Schweitzer, Feldmann, González-Barba and Vega, 2002 ENT#091;Campanian-Maastrichtian, Baja California SurENT#093;.
Costacopluma † Collins and Morris, 1975
Costacopluma bishopi † Vega and Feldmann, 1992 ENT#091;Coniacian, GuerreroENT#093; (Fraaije et al., 2006; Martínez-Díaz et al., 2016; Vega et al., 2016).
Costacopluma grayi † Feldmann and Portell, 2007 ENT#091;late Maastrichtian, CoahuilaENT#093; (Martínez-Díaz et al., 2016; Vega et al., 2016).
Costacopluma mexicana † Vega and Perrillat, 1989 ENT#091;late Campanian to early Maastrichtian of San Luis Potosí, Coahuila and Nuevo LeónENT#093; (Vega et al., 1995a; Martínez-Díaz et al., 2016; Vega et al., 2016) (Fig. 13F).
Superfamily CARPILIOIDEA Ortmann, 1893
Palaeoxanthopsidae † Schweitzer, 2003
Palaeoxanthopsis † Beurlen, 1958b
Palaeoxanthopsis meyapaquensis † (Vega, Feldmann, García-Barrera, Filkorn, Pimentel and Avendaño, 2001b, as Paraxanthopsis meyapaquensis) ENT#091;early Maastrichtian, ChiapasENT#093;.
Paraverrucoides † Schweitzer, 2003
Paraverrucoides alabamensis † (Rathbun, 1935b, as Xanthilites alabamensis) ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Vega et al., 2007a; Martínez-Díaz et al., 2017).
Verrucoides † Vega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a
Verrucoides stenohedra † Vega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a ENT#091;early Eocene (Ypresian), ChiapasENT#093; (Vega et al., 2008).
Tumidocarcinidae † Schweitzer, 2005a
Lobonotus † A. Milne-Edwards, 1863
Lobonotus mexicanus † Rathbun, 1930 ENT#091;Eocene, Baja California SurENT#093; (Schweitzer et al., 2002; 2006b; 2007a).
Zanthopsidae † Vía Boada, 1959
Neozanthopsis † Schweitzer, 2003
Neozanthopsis americanus † (Rathbun, 1928, as Harpactocarcinus americanus) ENT#091;?middle Eocene, Baja California SurENT#093; (Vega et al., 2006b).
Superfamily GONEPLACOIDEA MacLeay, 1838
Euryplacidae Stimpson, 1871
Orbitoplax † Tucker and Feldmann, 1990
Orbitoplax nandachare † (Vega et al., 2001a, as Stoaplax nandachare) ENT#091;Ypresian, ChiapasENT#093; (Vega et al., 2008).
Goneplacidae MacLeay, 1838
Amydrocarcinus † Schweitzer, Feldmann, González-Barba and Vega, 2002
Amydrocarcinus dantei † Schweitzer, Feldmann, González-Barba and Vega, 2002 ENT#091;middle Eocene (Bartonian), Baja California SurENT#093; (Schweitzer et al., 2007a).
Litocheiridae Števčić, 2005
Paracorallicarcinus † Tessier, Beschin, Bussulini and De Angeli, 1999
Paracorallicarcinus tricarinatus † Schweitzer, Feldmann, González-Barba, and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.
Mathildellidae Karasawa and Kato, 2003
Mathildellidae indet. ENT#091;late Campanian, CoahuilaENT#093; (Vega et al., 2016).
Tehuacana † Stenzel, 1944a
Tehuacana americana † (Rathbun, 1935, as Dromilites americana; = Tehuacana tehuacana Stenzel, 1944a) ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Vega et al., 2007a; 2008; Armstrong et al., 2009; Martínez-Díaz et al., 2017).
Tehuacana schweitzerae † Vega, Nyborg, Coutiño, and Hernández-Monzón, 2008 ENT#091;Eocene (Lutetian), ChiapasENT#093;.
Superfamily HEXAPODOIDEA Miers, 1886
Hexapodidae Miers, 1886
Palaeopinnixa † Vía Boada, 1966
Palaeopinnixa perornata † Collins and Morris, 1976 ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).
Superfamily PILUMNOIDEA Samouelle, 1819
Pilumnidae Samouelle, 1819
Eumedoninae Dana, 1853
Santeella † Blow and Manning, 1996
Santeella lillyae † Blow and Manning, 1996 ENT#091;early Eocene (Ypresian), ChiapasENT#093; (Vega et al., 2008).
Superfamily TRAPEZIOIDEA Miers, 1886
Trapeziidae Miers, 1886
Archaeotetra † Schweitzer, 2005b
Archaeotetra inornata † Schweitzer, 2005b ENT#091;middle Eocene, Baja California SurENT#093;.
Superfamily XANTHOIDEA MacLeay, 1838
Xanthoidea indet. ENT#091;middle Eocene, Baja California SurENT#093; (Schweitzer et al., 2006b).
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Eurytium Stimpson, 1862
Eurytium sp. ENT#091;early Miocene, Chiapas; Pliocene, VeracruzENT#093; (Vega et al., 2009).
Panopeus H. Milne Edwards, 1834
Panopeus veintensis † Vega, Nyborg, Coutiño and Hernández-Monzón, 2008 ENT#091;early Eocene (Ypresian), ChiapasENT#093;.
Xanthidae MacLeay, 1838
Xanthidae incertae sedis
?Haydnella † Müller, 1984
?Haydnella sp. cf. H. steiningeri † Müller, 1984 ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).
Megaxantho † Vega, Feldmann, Garcia-Barrera, Filkorn, Pimentel and Avendaño, 2001b
Megaxantho zoque † Vega, Feldmann, Garcia-Barrera, Filkorn, Pimentel and Avendaño, 2001b ENT#091;Maastrichtian, ChiapasENT#093; (Dietl and Vega, 2008).
Subsection THORACOTREMATA Guinot, 1977
Superfamily ?PINNOTHEROIDEA De Haan, 1833
?Pinnotheridae De Haan, 1833
?Pinnotherinae De Haan, 1833
Viapinnixa † Schweitzer and Feldmann, 2001
Viapinnixa alvarezi † Vega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a ENT#091;early Eocene (Ypresian), ChiapasENT#093; (Vega et al., 2008; Armstrong et al., 2009).
Viapinnixa perrilliatae † Vega, Nyborg, Fraaye and Espinosa, 2007a ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Armstrong et al., 2009) (Fig. 13K).
Superfamily GRAPSOIDEA MacLeay, 1838
Sesarmidae Dana, 1851c
Sesarmidae indet. ENT#091;Miocene (Aquitanian), ChiapasENT#093; (Serrano-Sánchez et al., 2016).
Brachyura incertae sedis
?Xanthosia Bell, 1863
? Xanthosia zoquiapensis † Fraaije, Vega, Van Bakel and Garibay-Romero, 2006 ENT#091;Campanian, GuerreroENT#093;.
Roemerus † Bishop, 1983b
Roemerus robustus † Bishop, 1983b ENT#091;Albian, ChiapasENT#093; (Vega et al., 2006a).
SOUTHERN AND CENTRAL FLORIDA, USA
Infraorder ANOMURA MacLeay, 1838
Superfamily GALATHEOIDEA Samouelle, 1819
Porcellanidae Haworth, 1825
Petrolisthes Stimpson, 1858
Petrolisthes myakkensis † Bishop and Portell, 1989 ENT#091;late Pliocene, Charlotte CountyENT#093; (Fig. 14A).
Fossil Anomura and Brachyura from southern and central Florida. A, Anomura: Galatheoidea: Porcellanidae: Petrolisthes myakkensisBishop and Portell, 1989, dorsal view of carapace of holotype UF 8678 adhered toHeliaster microbrachius (sun star), late Pliocene of El Jobean, Charlotte County. B, C, Brachyura: Raninoidea: Raninidae: Lophoranina sp. cf. L. georgiana(Rathbun, 1935), interior and lateral views of dorsal carapace of UF 67098, late Eocene of Ocala, Marion County. D, E, Eubrachyura: Calappoidea: Calappidae: Calappilia brooksi Ross and Scolaro, 1964, dorsal (D) and frontal (E) views of UF 13349, late Eocene of Newberry, Alachua County. F, Leucosioidea: Leucosiidae:Persephona mediterranea(Herbst, 1794), dorsal view of carapace and interior and exterior views of associated left cheliped UF 104495, early-middle Pleistocene of Fort Drum, Okeechobee County. G, H, Parthenopoidea: Parthenopidae: Platylambrus charlottensis (Rathbun, 1935), external and internal views of right propodus UF 40355, Arcadia, De Soto County. I, Majoidea: Mithracidae: Damithrax sp. cf. D. pleuracanthus (Stimpson, 1871), dorsal view of carapace of holotype UF 29057, Pliocene-Pleistocene of Sarasota, Sarasota County. Photos courtesy of Sean Roberts (FLMNH). All specimens dry, uncoated.
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Coenobita Latreille, 1829
Coenobitid indet. (crab habitational traces in neritid snail) ENT#091;early Pleistocene, Hendry CountyENT#093; (Vermeij and Portell, 2013).
Petrochirus Stimpson, 1858
Petrochirus bouvieri † Rathbun, 1918 ENT#091;early Pleistocene, Glades CountyENT#093; (Rathbun, 1935b).
Petrochirus diogenes (Linnaeus, 1758) ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Ranininae De Haan, 1839
Lophoranina † Fabiani, 1910
Lophoranina sp. cf. L. georgiana † (Rathbun, 1935b) ENT#091;late Eocene of Alachua, Marion, and Lafayette CountiesENT#093; (Portell, 2004) (Fig. 14B, C).
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Epialtidae MacLeay, 1838
Pisinae Dana, 1851b
Libinia Leach, 1815
Libinia sp. ENT#091;early-middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Mithracidae MacLeay, 1838
Damithrax Windsor and Felder, 2014
Damithrax sp. cf. D. pleuracanthus (Stimpson, 1871) ENT#091;late Pliocene to early Pleistocene, Sarasota CountyENT#093; (Klompmaker et al., 2015) (Fig. 14I).
Stenocionops Desmarest, 1823
Stenocionops suwanneeana † Rathbun, 1935b ENT#091;late Eocene, Suwannee CountyENT#093;.
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus sp. ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa ocalanus † (Ross, Lewis and Scolaro, 1964) (as Aparnocondylus) ENT#091;late Eocene (Priabonian) of Alachua, Marion, and Lafayette CountiesENT#093; (Portell, 2004; Rumsey et al., 2016).
Calappa robertsi † Ross, Lewis and Scolaro, 1964 ENT#091;late Eocene of Alachua, Marion, and Lafayette CountiesENT#093; (Portell, 2004).
Calappa spp. ENT#091;Plio-Pleistocene, central and southern peninsular FloridaENT#093; (Portell and Agnew, 2004).
Calappilia † A. Milne-Edwards, 1873
Calappilia brooksi † Ross and Scolaro, 1964 ENT#091;late Eocene of Alachua, Marion, and Lafayette CountiesENT#093; (Portell, 2004) (Fig. 14D, E).
Calappilia calculosa † Rumsey, Klompmaker and Portell, 2016 ENT#091;late Eocene-early Oligocene, Suwannee CountyENT#093;.
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Persephona Leach, 1817
Persephona mediterranea (Herbst, 1794) ENT#091;early-middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004) (Fig. 14F).
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Platylambrus Stimpson, 1871
Platylambrus charlottensis † (Rathbun, 1935b) ENT#091;early Pleistocene, Charlotte CountyENT#093; (Fig. 14G, H).
Superfamily CANCROIDEA Latreille, 1802
Cancridae Latreille, 1802
Cancrinae Latreille, 1802
Cancer Latreille, 1802
Cancer irroratus Say, 1817 ENT#091;late Pleistocene-Holocene, Miami-Dade CountyENT#093; (Rathbun, 1935b).
Superfamily CHEIRAGONOIDEA Ortmann, 1893
Cheiragonidae Ortmann, 1893
Montezumella † Rathbun, 1930
Montezumella microporosa † Portell and Collins, 2002 ENT#091;late Eocene, Alachua CountyENT#093; (Fig. 15F).
Fossil Eubrachyura from southern and central Florida. A, Carpilioidea: Carpiliidae: Ocalina floridana Rathbun, 1929, ventral view of UF 32725, late Eocene of Newberry, Alachua County; B, C, dorsal (B) and frontal (C) views of UF 105902, Newberry, Alachua County. D, E, Palaeocarpilius brodkorbi Lewis and Ross, 1965, dorsal (D) and frontal (E) views of UF 114368, late Eocene of Newberry, Alachua County. F, Cheiragonoidea: Cheiragonidae: Montezuemella microporosa Portell and Collins, 2002, dorsal view of holotype, UF 107150, late Eocene of Newberry, Alachua County. G, H, Eriphioidea: Menippidae: Menippe mercenaria (Say, 1818), dorsal (H) and frontal (G) views of carapace of UF 24668, late Pleistocene of Oldsmar, Pinellas County. I, Ocypodoidea: Ocypodidae: Ocypode quadrata (Fabricius, 1787), dorsal view of carapace, hypotype, UF 47573, late Pleistocene-Holocene of Satellite Beach, Brevard County. Photos courtesy of Sean Roberts (FLMNH). All specimens dry, uncoated.
Superfamily PORTUNOIDEA Rafinesque, 1815
Ovalipidae Spiridonov et al., 2014
Ovalipes Rathbun, 1898
Ovalipes stephensoniWilliams, 1976 ENT#091;early-middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Portunidae Rafinesque, 1815
Portuninae Rafinesque, 1815
Achelous De Haan, 1833
Achelous depressifrons (Stimpson, 1859) ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Achelous gibbesii (Stimpson, 1859) ENT#091;early-middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Achelous spinimanus (Latreille, 1819) ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004)
Portunus Weber, 1795
?Portunus sp. ENT#091;generic identification highly doubtful) ENT#091;middle Eocene, Citrus CountyENT#093; (Ivany et al., 1990; Portell, 2004).
Superfamily CARPILIOIDEA Ortmann, 1893
Carpiliidae Ortmann, 1893
Ocalina † Rathbun, 1929
Ocalina floridana † Rathbun, 1929 (type) ENT#091;late Eocene of Alachua, Levy, and Marion CountiesENT#093; (Rathbun, 1935b) (Fig. 15A-C).
Paraocalina † Beschin, Busulini, De Angeli and Tessier, 2007
Paraocalina brodkorbi † (Lewis and Ross, 1965) (as Palaeocarpilius brodkorbi) ENT#091;late Eocene, Alachua CountyENT#093; (Fig. 15D, E).
Superfamily ERIPHIOIDEA MacLeay, 1838
Menippidae Ortmann, 1893
Menippe De Hann, 1833
Menippe mercenaria (Say, 1818) ENT#091;late Pleistocene, Pinellas CountyENT#093; (Portell and Schindler, 1991) (Fig. 15G, H).
Menippe nodifronsStimpson, 1859 ENT#091;early Pleistocene, Glades CountyENT#093; (Rathbun, 1935b; Portell, 2004).
Superfamily PILUMNOIDEA Samouelle, 1819
Pilumnidae Samouelle, 1819
Pilumninae Samouelle, 1819
Pilumnus Leach, 1816
Pilumnus sp. ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Subsection THORACOTREMATA Guinot, 1977
Superfamily OCYPODOIDEA Rafinesque, 1815
Ocypodidae Rafinesque, 1815
Ocypode Weber, 1795
Ocypode quadrata (Fabricius, 1787) ENT#091;late Pleistocene-Holocene, Brevard CountyENT#093; (Rathbun, 1935b; Portell et al., 2003).
Uca Leach, 1814
Uca sp. ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Superfamily PINNOTHEROIDEA De Haan, 1833
Pinnotheridae De Haan, 1833
Pinnothereliinae Alcock, 1900a
Pinnixa White, 1846
Pinnixa sp. ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).
Caribbean Islands & Bermuda
ANGUILLA
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa earlei † Withers, 1924a ENT#091;early Miocene, Cartouche BayENT#093; (reported inWithers, 1924a, as late Oligocene; see also Collins et al. 2009c).
Calappa sp. (claw only) ENT#091;early Miocene, Cathedral CaveENT#093; (Collins et al., 2009c).
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Parthenope Weber, 1795
Parthenope sp. (claw fragment) ENT#091;early Miocene, Cartouche BayENT#093; (as late Oligocene inWithers, 1924a; see also Collins et al., 2009c).
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Scylla De Haan, 1833
Scylla costata † Rathbun, 1919 (claws only) ENT#091;early Miocene, West side of Road BayENT#093; (as late Oligocene inWithers, 1924a).
Podophthalminae Dana, 1851d
Psygmophthalmus † Schweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c
Psygmophthalmus bifurcatus † Collins in Collins et al., 2009c ENT#091;early Miocene, Betty Hill QuarryENT#093;.
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Panopeus H. Milne Edwards, 1834
Panopeus sp. (claw fragment) ENT#091;early Miocene, Cartouche BayENT#093; (reported inWithers, 1924a as late Oligocene; see also Collins et al. 2009c).
Brachyura incertae sedis
Brachyuran indet. (‘Lyreidus fastigatus’, claw fragment) ENT#091;Miocene, Crocus BayENT#093; (Feldmann, 1992) ENT#091;Note 1ENT#093;.
Note 1. A partial crab merus from Anguilla, initially described as representing the raninoid Lyreidus fastigatusRathbun, 1919, was later removed from the genus by Feldmann (1992) based on the dissimilar nature of the merus to Lyreidus. Due to the poor and fragmentary nature of the material, it cannot be assigned to a particular group with certainty, thus we consider it here as ‘Brachyura incertae sedis’.
ANTIGUA
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Coenobita Latreille, 1829
Coenobita sp. cf. C. clypeatus (Fabricius, 1787) ENT#091;late Holocene, Burma QuarryENT#093; (Luque, 2017) ) ENT#091;Note 1ENT#093;.
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Calappidae De Haan, 1833
Mursia Leach in Desmarest, 1823
Mursia granulosa †Collins and Donovan, 2002 ENT#091;late Oligocene, locality unknownENT#093; (Collins and Donovan, 2002, p. 145).
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes summus † Collins and Donovan, 1995 ENT#091;late Oligocene, Nonsuch BayENT#093; (Fig. 16A).
Fossil Anomura and Brachyura from the Neogene of the Caribbean. Antigua: A, Brachyura: Portunoidea: Portunidae: Necronectes summus Collins and Donovan, 1995, external view of chelipeds of UF 242593 embedded in limestone, Free Town, parish of Saint Philip. Curaçao: B, Brachyura: Raninoidea: Raninidae: Ranilia constricta(Milne-Edwards, 1880b), dorsal view of carapace of UF 227321, Saint Michiel. Jamaica: C, Anomura: Paguroidea: Diogenidae: Petrochirus bahamensis (Herbst, 1791), external view of right fixed finger and dactylus of UF 273849, Fort, parish of St. Thomas. D, Brachyura: Calappoidea: Calappidae: Calappa springeriRathbun, 1931, external view of right carpus, propodus, and dactylus of UF 273851, Fort, parish of St. Thomas. E, Dairoidea; Dairidae: Daira vulgaris Portell and Collins, 2004, dorsal view of carapace, holotype, UF 68349, Duncans, parish of Trelawny. F, Xanthoidea: Panopeidae: Lophopanopeus corallinus Portell and Collins, 2004, dorsal view of carapace, holotype, UF 106702, Duncans, parish of Trelawny. Puerto Rico: G, Portunoidea: Portunidae: Necronectes collinsi Schweitzer et al., 2006c, dorsal view of carapace of UF 231451, San Sebastian. Photos courtesy of Sean Roberts (FLMNH). All specimens dry, uncoated.
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Cardisoma Latreille, 1828
Cardisoma sp. cf. C. guanhumi Latreille, 1828 ENT#091;late Holocene, Burma QuarryENT#093; (Luque, 2017) ENT#091;Note 1ENT#093;.
GecarcinusLeach, 1814 ENT#091;in Leach, 1813-1815ENT#093;
Gecarcinus sp. aff. G. lateralis (Freminville, 1835) ENT#091;late Holocene, Burma QuarryENT#093; (Luque, 2017) ENT#091;Note 1ENT#093;.
Note 1. To date, these represent the first records of fossil anomurans and thoracotreme brachyurans from the island, and they are among the only fossil remains of Coenobita, Gecarcinus, and Cardisoma worldwide (Luque, 2017).
ARUBA
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Calappidae De Haan, 1833
Mursia Leach in Desmarest, 1823
Mursia creutzbergi † Collins and Donovan, 2004 ENT#091;Miocene-Pliocene, Rooi TakiENT#093;.
BAHAMAS
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
GecarcinusLeach, 1814 ENT#091;in Leach, 1813-1815ENT#093;
Gecarcinus sp. ENT#091;late Holocene, San SalvadorENT#093; (Locatelli, 2013).
BARBADOS
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Epialtidae MacLeay, 1838
Pisinae Dana, 1851b
Herbstia H. Milne Edwards, 1834
Herbstia exserta † Collins and Morris, 1976 ENT#091;middle to late Pleistocene, Coral RockENT#093;.
Mithracidae MacLeay, 1838
Maguimithrax Klompmaker et al., 2015
Maguimithrax spinosissimus (Lamarck, 1818) ENT#091;middle to late Pleistocene, HighgateENT#093; (Collins and Morris, 1976).
Mithrax Desmarest, 1823
Mithrax hemphilliRathbun, 1892 ENT#091;middle to late Pleistocene, HighgateENT#093; (Collins and Morris, 1976).
Mithrax hispidus (Herbst, 1790) ENT#091;as Mithrax caribbaeusRathbun, 1920aENT#093; ENT#091;middle to late Pleistocene, GibbonsENT#093; (Collins and Morris, 1976).
Mithrax aculeatus (Herbst, 1790) (as Mithrax verrucosus in H. Milne Edwards, 1832) ENT#091;middle to late Pleistocene of Gibbons, Clapham and GarrisonENT#093; (Collins and Morris, 1976).
Teleophrys Stimpson, 1860
Teleophrys ruber (Stimpson, 1871, as Mithraculus ruber) ENT#091;middle to late Pleistocene, GibbonsENT#093; (Collins and Morris, 1976).
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Atoportuninae Števčić, 2005
Laleonectes Manning and Chace, 1990
Laleonectes vocans (A. Milne-Edwards, 1878) ENT#091;middle to late Pleistocene, Coral RockENT#093;.
Portuninae Rafinesque, 1815
Achelous De Haan, 1833
Achelous gibbesii (Stimpson, 1862) ENT#091;middle to late Pleistocene, Coral RockENT#093;.
Superfamily CARPILIOIDEA Ortmann, 1893
Carpiliidae Ortmann, 1893
Carpilius Desmarest, 1823
Carpilius corallinus (Herbst, 1783) ENT#091;middle to late Pleistocene, Coral RockENT#093; (in Collins et al., 2009c).
Superfamily GONEPLACOIDEA MacLeay, 1838
Chasmocarcinidae Serène, 1964
Chasmocarcininae Serène, 1964
Falconoplax † Van Straelen, 1933b
Falconoplax bicarinella † Collins and Morris, 1976 ENT#091;early-middle Eocene Scotland Beds, SpaENT#093;.
Superfamily HEXAPODOIDEA Miers, 1886
Hexapodidae Miers, 1886
Palaeopinnixa † Vía Boada, 1966
Palaeopinnixa perornata † Collins and Morris, 1976 (type) ENT#091;early-middle Eocene Scotland Beds, SpaENT#093;.
Superfamily PILUMNOIDEA Samouelle, 1819
Pilumnidae Samouelle, 1819
Pilumninae Samouelle, 1819
Pilumnus Leach, 1816
Pilumnus sp. (claw fragment) ENT#091;Pleistocene, ClaphamENT#093; (Collins and Morris, 1976).
Superfamily XANTHOIDEA MacLeay, 1838
Xanthidae MacLeay, 1838
Actaeinae Alcock, 1898
Paractaea Guinot, 1969
Paractaea nodosa (Stimpson, 1860) ENT#091;Pleistocene, Coral RockENT#093; (as Actaea rufopunctata inCollins and Morris, 1976).
BERMUDA
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Cardisoma Latreille, 1828
Cardisoma sp. cf. C. guanhumi Latreille, 1828 ENT#091;late Pleistocene, Crystal CaveENT#093; (Luque, 2017) ENT#091;Note 1ENT#093;.
Gecarcinus Leach, 1814 ENT#091;in Leach, 1813-1815ENT#093;
Gecarcinus sp. ENT#091;late Pleistocene, Admiral CaveENT#093; (Luque, 2017; FLMNH online database ENT#091;Note 1ENT#093;.
Note 1. To our knowledge, these represent the first records of fossil decapods from Bermuda (Luque, 2017). The generic affinities of the several hundreds of Bermudan ‘Gecarcinus’ sp. fossil remains in the FLMNH Invertebrate Paleontology Collections are still to be confirmed, especially due to the potential occurrence of other non-congeneric terrestrial and semi-terrestrial crabs (J. Luque and R.W. Portell, pers. obs.).
BONAIRE
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Superfamily CALAPPOIDEA De Haan, 1833
Calappidae De Haan, 1833
Calappilia † A. Milne-Edwards, 1873
Calappilia bonairensis † Van Straelen, 1933a ENT#091;middle to late Eocene (Lutetian-Priabonian), southwest of Seroe MontagneENT#093;.
Superfamily CHEIRAGONOIDEA Ortmann, 1893
Cheiragonidae Ortmann, 1893
Montezumella † Rathbun, 1930
Montezumella rutteni † Van Straelen, 1933a ENT#091;middle to late Eocene (Lutetian-Priabonian), southwest of Seroe MontagneENT#093; (also reported inCollins and Donovan, 2005).
Superfamily CARPILIOIDEA Ortmann, 1893
Carpiliidae Ortmann, 1893
Ocalina † Rathbun, 1929
Ocalina sublevis † Collins and Donovan, 2005 ENT#091;middle to late Eocene (Lutetian-Priabonian), southwest of Seroe MontagneENT#093;.
CUBA
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
Paguridae Latreille, 1802
Paguridae indet. ENT#091;late Pleistocene, US Guantanamo Bay Naval StationENT#093; (Collins et al., 2009c).
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Ranininae De Haan, 1839
Vegaranina † Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a
Vegaranina precocia † (Feldmann, Vega, Tucker, García-Barrera and Avendaño 1996, as Lophoranina precocious) (type) ENT#091;Maastrichtian, Cienfuegos ProvinceENT#093; (Varela and Rojas-Consuegra, 2009).
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides sp. ENT#091;early Miocene, Matanzas ProvinceENT#093; (in Varela and Rojas-Consuegra, 2011b).
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Epialtidae MacLeay, 1838
Pisinae Dana, 1851b
Libinia Leach, 1815
Libinia sp. ENT#091;?Oligocene-Miocene, locality unknownENT#093; (Varela, 2013).
Mithracidae MacLeay, 1838
Mithrax Desmarest, 1823
Mithrax hispidus (Herbst, 1790) ENT#091;as M. caribbaeus inRathbun, 1920a) ENT#091;Pliocene-Pleistocene, La HabanaENT#093; (see also Peñalver et al., 1997).
?Mithrax sp. ENT#091;late Pliocene, Matanzas Province and Sancti Spíritus ProvincesENT#093; (Varela and Rojas Consuegra, 2009; Varela, 2013).
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus sp. ENT#091;early Miocene, Matanzas ProvinceENT#093; (in Varela and Rojas-Consuegra, 2011b).
Eriosachila † Blow and Manning, 1996
Eriosachila cubaensis † Varela and Rojas-Consuegra, 2011a ENT#091;early Miocene, Matanzas ProvinceENT#093;.
Calappidae De Haan, 1833
Calappidae indet. ENT#091;late Pleistocene, US Guantanamo Bay Naval StationENT#093; (Collins et al., 2009c).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Persephona Leach, 1817
Persephona sp. (merus fragment) ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (in Varela and Rojas-Consuegra, 2009).
Iliacantha Stimpson, 1871
Iliacantha sp. cf. I. liodactylusRathbun, 1898 ENT#091;early Miocene, Matanzas ProvinceENT#093; (inVarela and Rojas-Consuegra, 2011b).
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Spinolambrus Tan and Ng, 2007
Spinolambrus lazaroi † Varela, 2013 ENT#091;early Miocene, Matanzas ProvinceENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes collinsi † Schweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c ENT#091;early Miocene, Matanzas ProvinceENT#093; (Varela and Rojas-Consuegra, 2011b).
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphylax domingensis † (Rathbun, 1919) ENT#091;early Miocene, Matanzas ProvinceENT#093; (Schweitzer et al., 2006c; Varela and Rojas-Consuegra, 2011c).
Paraeuphylax † Varela and Schweitzer, 2011
Paraeuphylax cubaensis † (type) Varela and Schweitzer, 2011 ENT#091;early Miocene, Matanzas ProvinceENT#093;.
Portuninae Rafinesque, 1815
Arenaeus Dana, 1851d
Arenaeus sp. ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (Varela, 2013).
Callinectes Stimpson, 1862
Callinectes sp. ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (Varela, 2013).
Portunus Weber, 1795
Portunus oblongus † Rathbun, 1920b ENT#091;early Miocene, Matanzas ProvinceENT#093; (Varela and Rojas-Consuegra, 2009).
Portunus sp. (claw fragments) ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (Schweitzer et al., 2006c; Varela and Rojas-Consuegra, 2011c).
Superfamily ERIPHIOIDEA MacLeay, 1838
Eriphiidae MacLeay, 1838
Eriphia Latreille, 1817
Eriphia sp. ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (Varela, 2013).
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeidae indet. ENT#091;late Pleistocene, US Guantanamo Bay Naval StationENT#093; (Collins et al., 2009c).
Panopeinae Ortmann, 1893
Eurytium Stimpson, 1859
Eurytium sp. ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (Varela, 2013).
Panopeus H. Milne Edwards, 1834
Panopeus sp. ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (Varela and Rojas-Consuegra, 2009; Varela and Rojas-Consuegra, 2011c).
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Gecarcinus Leach, 1814 ENT#091;in Leach, 1813-1815ENT#093;
Gecarcinus ruricola (Linnaeus, 1758) ENT#091;late Pleistocene, Matanzas ProvinceENT#093; (identified by Mary J. Rathbun inRichards, 1935).
CURAÇAO
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Notopodinae Serène and Umali, 1972
Ranilia H. Milne Edwards, 1837
Ranilia constricta (A. Milne-Edwards, 1880b) ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014) (Fig. 16B).
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides lamarcki A. Milne-Edwards and Bouvier, 1923 ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014).
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Mithracidae MacLeay, 1838
Maguimithrax Klompmaker et al., 2015
Maguimithrax spinosissimus (Lamarck, 1818) ENT#091;Pliocene, Salina Sint MichaelENT#093; (Stepp, 2014; Klompmaker et al., 2015).
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus lineatinus † Collins and Todd in Todd and Collins, 2005 ENT#091;Pliocene, Salina Saint MichielENT#093; (Stepp, 2014).
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa galloidesStimpson, 1859 ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Myropsis Stimpson, 1871
Myropsis quinquespinosaStimpson, 1871 ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014)
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Portuninae Rafinesque, 1815
Achelous De Haan, 1833
Achelous sebae (H. Milne Edwards, 1834) ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014).
DOMINICAN REPUBLIC
Infraorder ANOMURA MacLeay, 1838
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Dardanus Paul’son, 1875
Dardanus squamatus † Collins in Collins et al., 2009c ENT#091;late Miocene-early Pliocene, Río Gurabo and Río MaoENT#093;.
Petrochirus Stimpson, 1858
Petrochirus inequalis † Rathbun, 1919 ENT#091;Pleistocene, Rio Gurabo and Santo DomingoENT#093; (Collins et al., 2009c).
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus guraboensis † Collins in Collins et al., 2009c ENT#091;late Miocene, Rio Gurabo, Rio Cana and Rio MaoENT#093;.
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa flammea (Herbst, 1794) ENT#091;late Miocene, Rio Gurabo and Santo DomingoENT#093; (Collins et al., 2009c).
Calappa sp. ENT#091;late Miocene-early Pliocene, Rio GuraboENT#093; (Collins et al., 2009c).
Cryptosoma Brullé, 1839
Cryptosoma bairdii (Stimpson, 1862) ENT#091;late Miocene, Rio GuraboENT#093; (Collins et al., 2009c).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Persephona Leach, 1817
Persephona prepunctata † Rathbun, 1919 ENT#091;late Miocene, Rio Gurabo and Rio MaoENT#093;.
Iliacantha Stimpson, 1871
Iliacantha sp. ENT#091;early Miocene, Rio GuraboENT#093; (Collins et al., 2009c).
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Mesorhoea Stimpson, 1871
Mesorhoea mauryae † Rathbun, 1919 ENT#091;late Miocene-early Pliocene, Río Cana and Santo DomingoENT#093; (Collins et al., 2009c).
Platylambrus Stimpson, 1871
Platylambrus obscura † Rathbun, 1919 ENT#091;Miocene, Santo DomingoENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Scylla De Haan, 1833
Scylla costata † Rathbun, 1919 ENT#091;Miocene, valley of Yaque del Norte RiverENT#093; ENT#091;Note 1ENT#093;.
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphylax domingensis † (Rathbun, 1919, as Podophthalmus domingensis) ENT#091;Miocene, valley of Yaque del Norte RiverENT#093; ENT#091;Note 1ENT#093;.
Psygmophthalmus † Schweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c
Psygmophthalmus bifurcatus † Collins in Collins et al., 2009c ENT#091;age uncertain, Rio CanaENT#093;.
Sandomingia † Rathbun, 1919
Sandomingia yaquiensis † Rathbun, 1919 ENT#091;late Miocene-early Pliocene, Río Cana and Rio Yaque del NorteENT#093; (Collins et al., 2009c) ENT#091;Note 1ENT#093;.
Portuninae Rafinesque, 1815
Callinectes Stimpson, 1862
Callinectes declivis † Rathbun, 1918 (fingers only) ENT#091;late Miocene, Rio Gurabo and Rio Mao Santo DomingoENT#093; (inCollins et al., 2009c).
Portunus Weber, 1795
Portunus gabbi † Rathbun, 1919 ENT#091;Miocene, valley of Yaque del Norte RiverENT#093; ENT#091;Note 1ENT#093;.
Portunus oblongus † Rathbun, 1920b ENT#091;late Miocene-early Pliocene, Río CanaENT#093;.
Portunus tenuis † Rathbun, 1919 (type) (claws only) ENT#091;Miocene of Yaque Valley, Santo DomingoENT#093;.
Rathbunites † Schweitzer, Dworschak and Martin, 2011 (as Rathbunella in Collins et al., 2009c , name preoccupied)
Rathbunites pentaspinosa (Collins in Collins et al., 2009c) (type) ENT#091;late Miocene-early Pliocene, Río CanaENT#093;.
Superfamily CARPILIOIDEA Ortmann, 1893
Tumidocarcinidae † Schweitzer, 2005a
Lobonotus † A. Milne-Edwards, 1863
Lobonotus sculptus † A. Milne-Edwards, 1863 (as Archaeopilumnus caelatus inRathbun, 1919) ENT#091;late Miocene, Rio GuraboENT#093; (Rathbun, 1919; 1920b; Collins et al., 2009c; Ossó et al., 2014).
Superfamily PILUMNOIDEA Samouelle, 1819
Pilumnidae Samouelle, 1819
Pilumninae Samouelle, 1819
Pilumnus Leach, 1816
Pilumnus subequus † Rathbun, 1919 (claws only) ENT#091;Miocene of Yaque Valley, Santo DomingoENT#093; ENT#091;Note 1ENT#093;.
Pilumnus sp. ENT#091;late Miocene, Rio GuraboENT#093; (Collins et al., 2009c).
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Eurypanopeus A. Milne-Edwards, 1880a
Eurypanopeus sp. aff. E. crenatus H. Milne Edwards, 1834 ENT#091;age uncertain, Rio CanaENT#093; (Collins et al., 2009c).
Panopeus H. Milne Edwards, 1834
Panopeus sp. ENT#091;late Miocene, Rio GuraboENT#093; (Collins et al., 2009c).
Note 1. As previously noted by Collins et al. (2009c, p. 62), Rathbun (1919) reported a handful of fossil crabs ENT#091;Scylla costata, Portunus gabbi, Euphylax dominguensis (as Podophthalmus), and Sandomingia yaquiensisENT#093; from Haiti, but they originate from lower Miocene rocks exposed along the “Lower half of the valley of the Yaqui ENT#091;sicENT#093; del Norte River, in the northern part of Santo Domingo” in the Dominican Republic.
THE GRENADINES
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa sp. cf. C. springeri (Rathbun, 1931) ENT#091;middle Miocene, CarriacouENT#093; (Donovan et al., 2003; Collins et al., 2009c).
Superfamily ERIPHIOIDEA MacLeay, 1838
Platyxanthidae Guinot, 1977
Platyxanthus A. Milne-Edwards, 1863
Platyxanthus sp. ENT#091;middle Miocene, CarriacouENT#093; (Collins et al., 2009c).
HAITI
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Eubrachyuran indet. (claw) (as ?Zanthopsis indet. inRathbun, 1923a) ENT#091;Eocene, Artibonite departmentENT#093; ENT#091;Note 2ENT#093;.
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Mithracidae MacLeay, 1838
Mithrax Desmarest, 1823
?Mithrax sp. ENT#091;Pleistocene, Môle St. Nicolas in Nord-West departmentENT#093; (Rathbun, 1923a) ENT#091;Note 2ENT#093;.
Superfamily PARTHENOPOIDEA MacLeay, 1838
?Parthenopoid indet. (claw fragment) ENT#091;early Miocene, Centre departmentENT#093; ENT#091;Note 2ENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Portuninae Rafinesque, 1815
Portunus Weber, 1795
Portunus haitensis † Rathbun, 1923a ENT#091;early Miocene, Centre departmentENT#093; ENT#091;Note 1ENT#093;.
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Panopeus H. Milne Edwards, 1834
?Panopeus sp. indet. (fingers only) ENT#091;early Miocene, Centre departmentENT#093; (Rathbun, 1923a) ENT#091;Note 2ENT#093;.
Note 1.Rathbun (1923a) assigned some fossil cheliped fragments from Haiti to Parthenope, Panopeus, and ‘Zanthopsis’ with uncertainty. Indeed, the generic and familial placements need to be re-examined.
Note 2. As previously noticed by Collins et al. (2009c, p. 62), a handful of fossil crabs reported by Rathbun (1919) as coming from Haiti are in fact from the Miocene of Dominican Republic.
JAMAICA
Infraorder ANOMURA MacLeay, 1838
Superfamily GALATHEOIDEA Samouelle, 1819
Porcellanidae Haworth, 1825
Petrolisthes Stimpson, 1858
Petrolisthes sp. ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Superfamily HIPPOIDEA Latreille, 1825
Albuneidae Stimpson. 1858
Albunea Weber, 1975
Albunea sp. ENT#091;late Pleistocene, parish of St. Ann) (Morris, 1993).
Superfamily PAGUROIDEA Latreille, 1802
Diogenidae Ortmann, 1892
Dardanus Paul’son, 1875
Dardanus portmorantensis † Collins and Donovan, 2012 ENT#091;Pleistocene, parish of St. ThomasENT#093;.
Petrochirus Stimpson, 1858
Petrochirus bahamensis (Herbst, 1791) ENT#091;late Pliocene to late Pleistocene, parish of St. ThomasENT#093; (Collins and Portell, 1998; Collins et al., 2009a; Collins and Donovan, 2012) (Fig. 16C).
Petrochirus sp. ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Paguristes Dana, 1851c
Paguristes sp. cf. P. lymanni A. Milne-Edwards and Bouvier, 1893 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Paguristes sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Infraorder BRACHYURA Latreille, 1802
Superfamily HOMOLODROMIOIDEA Alcock, 1900b
Goniodromitidae † Beurlen, 1932
Trechmannius † Collins and Donovan, 2006
Trechmannius circularis † Collins and Donovan, 2006 (type) ENT#091;early Paleocene, parish of PortlandENT#093;.
Superfamily DROMIOIDEA De Haan, 1833
Dynomenidae Ortmann, 1892
Dynomene Desmarest, 1823
Dynomene variabilis † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Kromtitis † Müller, 1984
Kromtitis spinulata † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Superfamily PALAEOCORYSTOIDEA † Lőrenthey in Lőrenthey and Beurlen, 1929
Palaeocorystidae † Lőrenthey in Lőrenthey and Beurlen, 1929
Cretacoranina † Mertin, 1941
Cretacoranina trechmanni † (Withers, 1927, as Ranina trechmanni) ENT#091;Maastrichtian, parish of St. JamesENT#093; (Morris, 1993).
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Raninoidinae Lőrenthey in Lőrenthey and Beurlen, 1929
Raninoides H. Milne Edwards, 1837
Raninoides louisianensisRathbun, 1933 ENT#091;Pleistocene, late Pleistocene, parish of St. ThomasENT#093; (Collins and Donovan, 1998).
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily MAJOIDEA Samouelle, 1819
Epialtidae MacLeay, 1838
Pisinae Dana, 1851b
Chlorilia Dana, 1851b
Chlorilia sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Libinia Leach, 1815
Libinia milnei (Collins and Donovan, 2012) ENT#091;Pleistocene, parish of PortlandENT#093;.
Pitho Bell, 1836
Pitho anisodon (von Martens, 1872) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 1997).
Pitho sp. (claw dactylus) ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Rochinia A. Milne-Edwards, 1875
Rochinia sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Mithracidae MacLeay, 1838
Damithrax Windsor and Felder, 2014
Damithrax unguis † (Portell and Collins, 2004, as Mithrax unguis) ENT#091;early Miocene, parish of TrelawnyENT#093; (Klompmaker et al., 2015).
Maguimithrax Klompmaker et al., 2015
Maguimithrax spinosissimus (Lamarck, 1818) ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Mithraculus White, 1847
Mithraculus forceps A. Milne-Edwards, 1875 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a) ENT#091;Note 1ENT#093;.
Mithraculus sp. aff. M. coryphe (Herbst, 1790) ENT#091;late Miocene, parish of St. ThomasENT#093; (Collins et al., 2010).
Mithraculus sp. cf. M. forceps A. Milne-Edwards, 1875 ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Mithrax Desmarest, 1823
Mithrax arawakum † Klompmaker et al., 2015 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Mithrax sp. cf. M. hispidus (Herbst, 1790) ENT#091;= M. caribbaeusRathbun, 1920aENT#093; ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Mithrax aculeatus (Herbst, 1782-1804) (= M. verrucosus H. Milne Edwards, 1832) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Nemausa A. Milne-Edwards, 1875
Nemausa acuticornis (Stimpson, 1871) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Nemausa donovani † (Portell and Collins, 2004, as Mithrax donovani) ENT#091;early Miocene, parish of TrelawnyENT#093;.
Nemausa windsorae † Klompmaker et al., 2015 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Teleophrys Stimpson, 1860
Teleophrys acornis † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Oregoniidae Garth, 1958
Hyas Leach, 1814 ENT#091;in Leach, 1813-1815ENT#093;
Hyas sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus praecox † Collins, Donovan and Dixon, 1997 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Hepatus sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa sp. cf. C. gallus (Herbst, 1803) ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Calappa springeriRathbun, 1931 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 1997) (Fig. 16D).
Calappa sp. aff. C. springeriRathbun, 1931 ENT#091;late Pliocene, parish of St. Thomas ENT#093; (Collins and Portell, 1998).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Persephona Leach, 1817
Persephona punctata (Linnaeus, 1758) ENT#091;late Pleistocene, parish of St. Thomas) (Collins et al., 1997).
Persephona sp. aff. P. punctata (Linnaeus, 1758) ENT#091;late Pliocene, parish of St. Ann) (Morris, 1993).
?Persephona sp. ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Uhlias Stimpson, 1871
Uhlias sp. cf. U. limbatusStimpson, 1871 ENT#091;late Pleistocene, parish of St. Ann) (Morris, 1993).
Leucosiidae incertae sedis
Duncanitrix † Schweitzer, Dworschak and Martin, 2011 ENT#091;=Duncania Portell and Collins, 2004 ENT#093;
Duncanitrix jamaicensis † (Portell and Collins, 2004) ENT#091;early Miocene, parish of TrelawnyENT#093;.
Superfamily PARTHENOPOIDEA MacLeay, 1838
Parthenopidae MacLeay, 1838
Parthenopinae MacLeay, 1838
Mesorhoea Stimpson, 1871
Mesorhoea sp. aff. M. sexspinosaStimpson, 1871 ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Platylambrus Stimpson, 1871
Platylambrus sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Superfamily CANCROIDEA Latreille, 1802
Cancridae Latreille, 1802
Cancrinae Latreille, 1802
Cancer Linnaeus, 1758
Cancer sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Superfamily PORTUNOIDEA Rafinesque, 1815
Carcineretidae † Beurlen, 1930
Carcineretes † Withers, 1922
Carcineretes woolacotti † Withers, 1922 ENT#091;Maastrichtian, parishes of Clarendon and St. JamesENT#093;.
Ovalipidae Spiridonov et al., 2014
Ovalipes Rathbun, 1898
Ovalipes sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Portunidae Rafinesque, 1815
Atoportuninae Števčić, 2005
Laleonectes Manning and Chace, 1990
Laleonectes vocans (A. Milne-Edwards, 1878) ENT#091;late Pleistocene, parish of St. ThomasENT#093;.
Podophthalminae Dana, 1851d
Euphylax Stimpson, 1862
Euphylax fortispinosus † Collins, Donovan, Lindsay and Simpson, 2001 ENT#091;early Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009c).
Portuninae Rafinesque, 1815
Achelous De Haan, 1833
Achelous sebae (H. Milne Edwards, 1834) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Callinectes Stimpson, 1862
Callinectes jamaicensis † Withers, 1924b (claw fragment only) ENT#091;middle Eocene (Lutetian), parish of HanoverENT#093; (Morris, 1993).
Callinectes sp. aff. C. sapidusRathbun, 1896 ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Callinectes sp. cf. C. toxodesOrdway, 1863 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 1997).
Portunus Weber, 1795
Portunus sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Pseudoachelous † Portell and Collins, 2004
Pseudoachelous schindleri † Portell and Collins, 2004 (type) ENT#091;early Miocene, parish of TrelawnyENT#093;.
Superfamily CARPILIOIDEA Ortmann, 1893
Carpiliidae Ortmann, 1893
Carpilius Desmarest, 1823
Carpilius corallinus (Herbst, 1783) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Ocalina † Rathbun, 1929
Ocalina haldixoni † Collins and Donovan, 2006 ENT#091;middle to late Eocene, parish of PortlandENT#093;.
Palaeoxanthopsidae † Schweitzer, 2003
?Palaeoxanthopsidae incertae sedis † (as Necrocarcinus sp. inMorris, 1993) ENT#091;?Maastrichian, unknownENT#093; ENT#091;Note 2ENT#093;.
Palaeoxanthopsis † Beurlen, 1958b
Palaeoxanthopsis sp. † (as ?Paranecrocarcinus sp. inMorris, 1993, fig. 1.6) ENT#091;?Maastrichian, unknownENT#093; ENT#091;Note 2ENT#093;.
Superfamily DAIROIDEA Serène, 1965
Dairidae Serène, 1965
Daira De Haan, 1833
Daira vulgaris † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093; (Fig. 16E)
Superfamily ERIPHIOIDEA MacLeay, 1838
Eriphiidae MacLeay, 1838
Eriphia Latreille, 1817
Eriphia gonagra xaymacaensis † Collins and Donovan, 1998 (type) ENT#091;late Pleistocene, parish of St. ThomasENT#093;.
Eriphia sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Superfamily PILUMNOIDEA Samouelle, 1819
Pilumnidae Samouelle, 1819
Pilumninae Samouelle, 1819
Pilumnus Leach, 1816
Pilumnus sp. aff. P. pannosusRathbun, 1898 ENT#091;late Pliocene, parish of St. ThomasENT#093;.
Pilumnus sp. aff. P. sayiRathbun, 1897a ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 1997).
Pilumnus sp. aff. P. spinossimusRathbun, 1898 ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Superfamily TRAPEZIOIDEA Miers, 1886
Trapeziidae Miers, 1886
Trapezia Latreille, 1828
Trapezia prisca † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Eurypanopeus A. Milne-Edwards, 1880a
Eurypanopeus abbreviatus (Stimpson, 1860) ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Eurypanopeus sp. cf. E. depressus (Smith, 1869b) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 1997; Collins and Donovan, 2012).
Eurypanopeus sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Eurytium Stimpson, 1859
Eurytium sp. cf. E. limosum (Say, 1818) ENT#091;late Pliocene to late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 1997; 2009c; Collins and Portell, 1998).
Hexapanopeus Rathbun, 1898
Hexapanopeus caribbaeus (Stimpson, 1871) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 1997).
Lophopanopeus Rathbun, 1898
Lophopanopeus corallinus † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093; (Fig. 16F).
Lophopanopeus toomeyorum † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Neopanope A. Milne-Edwards, 1880a
Neopanope sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Panopeus H. Milne Edwards, 1834
Panopeus herbstii H. Milne Edwards, 1834 (claw only) ENT#091;late Pliocene-late Pleistocene, parishes of St. Thomas and St. AnnENT#093; (Morris, 1993; Collins and Portell, 1998).
Panopeus nanus † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Panopeus rugosus A. Milne-Edwards, 1880b (claw only) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins and Donovan, 2012).
Tetraxanthus Rathbun, 1898
Tetraxanthus sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins et al., 2009) (listed in Collins and Portell, 1998 as aff. Eurypoda sp.).
Pseudorhombilidae Alcock, 1900a
Nanoplax Guinot, 1967
Nanoplax xanthiformis (A. Milne-Edwards, 1880b) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Xanthidae MacLeay, 1838
Actaeinae Alcock, 1898
Actaea De Haan, 1833
Actaea acantha (H. Milne Edwards, 1834) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Actaea bifronsRathbun, 1898 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Actaeops † Portell and Collins, 2004
Actaeops frontalis † Portell and Collins, 2004 (type) ENT#091;early Miocene, parish of TrelawnyENT#093;.
Chlorodiellinae Ng and Holthuis, 2007
Chlorodiella Rathbun, 1897b (= Chlorodius H. Milne Edwards, 1834 )
Chlorodiella occidentalis † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Xanthinae MacLeay, 1838
Leptodius A. Milne-Edwards, 1863
Leptodius granulatus † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Micropanope Stimpson, 1871
? Micropanope nuttingi (Rathbun, 1898) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).
Micropanope sp. cf. M. politaRathbun, 1893 ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993; Collins et al., 2001).
Micropanope sp. cf. M. spinipes A. Milne-Edwards, 1988a ENT#091;late Pliocene to late Pleistocene, parishes St. Thomas and St. AnnENT#093; (Morris, 1993; Collins and Portell, 1998).
Micropanope pulcherrima † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.
Micropanope sp. aff. M. truncatifronsRathbun, 1898 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).
Cycludius Dana, 1851 (= Phymodius A. Milne-Edwards, 1863 )
Cycludius sp. cf. C. maculatus (Stimpson, 1860) ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Xanthoidea incertae sedis
?Xanthilites rathbunae † Withers, 1924b ENT#091;middle Eocene (Lutetian), parish of HanoverENT#093; (Morris, 1993).
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Cardisoma Latreille, 1828
Cardisoma guanhumi Latreille, 1828 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins and Donovan, 1998; 2012).
Grapsidae MacLeay, 1838
Pachygrapsus Randall, 1840
Pachygrapsus sp. ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Sesarmidae Dana, 1851c
Sesarma Say, 1817
Sesarma sp. cf. S. cookeiHartnoll, 1971 (as S. primigenium inCollins, Mitchell and Donovan, 2009b) ENT#091;late Pleistocene, parish of St. ElizabethENT#093; (Donovan and Dixon, 1998; Baalbergen and Donovan, 2013; Donovan, 2016) ENT#091;Note 3ENT#093;.
Varunidae H. Milne Edwards, 1853
?Varuna H. Milne Edwards, 1830
?Varuna sp. ENT#091;middle Eocene, parish of St. JamesENT#093; (Morris, 1993).
Superfamily OCYPODOIDEA Rafinesque, 1815
Ocypodidae Rafinesque, 1815
Uca Leach, 1814
Uca sp. ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).
Note 1.Collins et al. (2009a) listed this taxon from the late Pliocene Bowden Formation in error.
Note 2. Two fragmentary dorsal carapaces from the late Cretaceous (stage unknown) were assigned by Morris (1993) to the necrocarcinoids Necrocarcinus and ?Paranecrocarcinus (Morris, 1993, fig. 1.5 and 1.6, respectively), but both Jamaican specimens lack diagnostic features of the families Necrocarcinidae and Paranecrocarcinidae. The overall configuration of the carapace regions and grooves, the absence of longitudinal rows of tubercles axially and along the branchial regions, the lack of coarse granulations, and the absence of postrostral slits, preclude placement among Necrocarcinus, ?Paranecrocarcinus, or any genus or family of Necrocarcinoidea (Schweitzer et al., 2016). The specimens seem better placed among the heterotremous Eubrachyura, particularly the family Palaeoxanthopsidae. The specimen referred to ‘?Paranecrocarcinus’ seems congeneric with PalaeoxanthopsisBeurlen, 1958b, and shares some overall similarities with P. tylotus from the late Maastrichtian of Puerto Rico (Schweitzer et al., 2008). The specimen reported as ‘Necrocarcinus’ also has an overall palaeoxanthopsid appearance, but its systematic affinities are yet to be determined.
Note 3. Cheliped remains of Sesarma sp. cf. S. cookei from the late Pleistocene of Jamaica were initially identified as Gecarcinus cf. ruricola (Collins in Donovan and Dixon, 1998), later as belonging to S. primigenium (Collins et al., 2009b), and more recently as Sesarma sp. cf. S. cookei (Baalbergen and Donovan, 2013; Donovan, 2016; Luque, 2017).
PUERTO RICO
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Ranininae De Haan, 1839
Vegaranina † Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a
Vegaranina precocia † (Feldmann, Vega, Tucker, García-Barrera and Avendaño 1996, as Lophoranina precocious) ENT#091;late Maastrichtian, Sabana GrandeENT#093; (Schweitzer et al., 2008).
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa pavimenta † Schweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c ENT#091;early Miocene, San SebastiánENT#093;.
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes collinsiSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c ENT#091;Oligocene-Miocene, San Sebastián and YaucoENT#093; (Schweitzer et al., 2008) (Fig. 16G).
Necronectes proavitus † (Rathbun, 1918) ENT#091;Miocene, uncertainENT#093; (Gordon, 1966; Schweitzer et al., 2006c).
Scylla De Haan, 1833
Scylla costata † Rathbun, 1919 (Miocene, San Sebastián and YaucoENT#093; (Schweitzer et al., 2008).
Podophthalminae Dana, 1851d
Psygmophthalmus † Schweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c
Psygmophthalmus lares † Schweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c (type) ENT#091;early Miocene, San SebastiánENT#093;.
Portuninae Rafinesque, 1815
Portunus Weber, 1795
Portunus oblongus † Rathbun, 1920b ENT#091;Miocene, uncertainENT#093; (Gordon, 1966; Schweitzer et al., 2006c).
Portunus yaucoensis † Schweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c ENT#091;early Oligocene (Rupelian), YaucoENT#093;.
Superfamily CARPILIOIDEA Ortmann, 1893
Palaeoxanthopsidae † Schweitzer, 2003
Palaeoxanthopsis † Beurlen, 1958b
Palaeoxanthopsis tylotus † Schweitzer, Velez-Juarbe, Martinez, Collmar Hull, Feldmann and Santos, 2008 ENT#091;late Maastrichtian, Sabana GrandeENT#093;.
Superfamily XANTHOIDEA MacLeay, 1838
Panopeidae Ortmann, 1893
Panopeinae Ortmann, 1893
Eurytium Stimpson, 1859
Eurytium granulosum † Schweitzer, Velez-Juarbe, Martinez, Hull, Feldmann and Santos, 2008 ENT#091;Miocene, near PonceENT#093; (Schweitzer et al., 2008).
Subsection THORACOTREMATA Guinot, 1977
Superfamily GRAPSOIDEA MacLeay, 1838
Gecarcinidae MacLeay, 1838
Cardisoma Latreille, 1828
Cardisoma guanhumiLatreille, 1828 ENT#091;Pleistocene, UtuadoENT#093; (Schweitzer et al., 2008).
SAINT BARTHÉLEMY (ST. BARTS)
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Eriosachila † Blow and Manning, 1996
Eriosachila bartholomaensis † (Rathbun, 1919, as Zanthopsis bartholomaensis) ENT#091;Eocene, St. BartholomewENT#093;.
SAINT MARTIN
Infraorder BRACHYURA Latreille, 1802
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Calappidae De Haan, 1833
Tutus † Collins in Collins et al., 2009c
Tutus granulosus † Collins in Collins et al., 2009c ENT#091;Miocene of Leeward Islands, Tintamare IslandENT#093;.
TRINIDAD
Infraorder BRACHYURA Latreille, 1802
Superfamily RANINOIDEA De Haan, 1839
Raninidae De Haan, 1839
Ranininae De Haan, 1839
Lophoranina † Fabiani, 1910
Lophoranina porifera † (Woodward, 1866) ENT#091;‘Tertiary’ indet, San FernandoENT#093;.
Section EUBRACHYURA Saint Laurent, 1980
Subsection HETEROTREMATA Guinot, 1977
Superfamily CALAPPOIDEA De Haan, 1833
Aethridae Dana, 1851d
Hepatus Latreille, 1802
Hepatus nodosus † Collins and Morris, 1976 ENT#091;middle Miocene, Manzanilla BayENT#093;.
Calappidae De Haan, 1833
Calappa Weber, 1795
Calappa cuspidata † (Guppy, 1909, as Ranina cuspidata) ENT#091;Oligocene, Tamana DistrictENT#093; (Pasini and Garassino, 2017).
Superfamily LEUCOSIOIDEA Samouelle, 1819
Leucosiidae Samouelle, 1819
Ebaliinae Stimpson, 1871
Persephona Leach, 1817
Persephona punctata (Linnaeus, 1758) ENT#091;Miocene-Pliocene, Manzanilla BayENT#093; (Collins and Morris, 1976).
Superfamily PORTUNOIDEA Rafinesque, 1815
Portunidae Rafinesque, 1815
Necronectinae Glaessner, 1928
Necronectes † A. Milne-Edwards, 1881
Necronectes proavitus † (Rathbun, 1918) ENT#091;middle Miocene, near Caparo Saca MantecaENT#093; (Collins and Morris, 1976).
Portuninae Rafinesque, 1815
Portunus Weber, 1795
Portunus oblongus † Rathbun, 1920b ENT#091;middle Miocene, Manzanilla Bay and MontserratENT#093; (Collins and Morris, 1976).
Superfamily HEXAPODOIDEA Miers, 1886
Hexapodidae Miers, 1886
Palaeopinnixa † Vía Boada, 1966
Palaeopinnixa intermedia † (Collins and Morris, 1976, as Thaumastoplax intermedia) ENT#091;middle Miocene, MontserratENT#093;
Palaeopinnixa perornata † Collins and Morris, 1976 (type) ENT#091;Miocene, MontserratENT#093;.
Final Remarks
This revision of the anomuran and brachyuran fossil record in the tropical Americas demonstrates a diversity and abundance previously unassessed, with 32 superfamilies, 69 families, 190 genera, and 415 species properly recognized (Fig. 1D, Chart 1). Most superfamilies and families in this checklist have pantropical distributions, although there is a considerable degree of endemism at the generic and specific levels, particularly during the early Cretaceous of northern South America and Mexico, e.g., Brazilomunida, Protaegla, Maurimia, Dynomenopsis, Araripecarcinus, Bellcarcinus, Colombicarcinus, Planocarcinus, Tepexicarcinus, and several new Cretaceous and Paleogene families, genera and species under study. However, the sampling bias in the tropics is still high, with most fossil records known from Cenozoic deposits through the continental and insular Caribbean. This is exemplified by countries like Jamaica (with 32 families, 65 genera, and 71 species) or Panama (with 24 families, 41 genera, and 47 species), that independently account for more fossil occurrences than much larger countries like Bolivia, Colombia, Ecuador and Peru all combined (Fig. 1D, Chart 1).
Despite this, several recent findings - particularly from northern South America - represent the oldest members of their genera, families, superfamilies, or even new lineages, and suggest a more complex phylogenetic scenarios than currently depicted (Luque et al., 2016). Furthermore, several of these new findings from challenge the paradigm of a high latitude origin for several groups, and extend their stratigraphic and paleogeographic ranges into the equatorial Neotropics, highlighting the role and importance of the tropical Americas in the origin and evolution of decapod crustaceans through time.
Acknowledgements
JL thanks Carlos Jaramillo (Smithsonian Tropical Research Institute, Panamá) and A. Richard Palmer (University of Alberta, Canada) for providing partial funds and facilities; Sylvain Charbonnier, Jocelyn Falconnet, and Peter Massicard (MNHN, Paris, France), and Rodney M. Feldmann (Kent State University, USA), for providing images of fossil specimens from the MNHN and USNM collections, respectively; Wade Jones (Kent State University, USA), Gale Bishop (Georgia Southern University, Georgia, USA), Stephen K. Donovan (Naturalis Biodiversity Center, Leiden, The Netherlands), Günter Schweigert (Staatliches Museum für Naturkunde, Stuttgart, Germany), Thomas Hegna (Western Illinois University, USA), and Gilberto Juárez Huarachi (Bolivia), for providing literature items. JL also thanks the Autoridad del Canal de Panamá (ACP) for providing laboratory facilities; the Dirección de Recursos Minerales de Panamá for facilitating collecting and export permits; Ricardo Perez for donating the Toyota vehicles used for fieldwork in Panama; Álex Ossó (Tarragona, Catalonia) for valuable discussions; Austin J.W. Hendy (Natural History Museum of Los Angeles County, USA), Kecia Kerr (University of Alberta, Canada), Jorge W. Moreno (Universidad del Norte, Barranquilla, Colombia), several friends and colleagues from the Smithsonian Tropical Research Institute, Center for Tropical Paleoecology and Archaeology (STRI CTPA), and the Colombian Geological Survey for field assistance; Daniel Levin (Dept. of Paleobiology, Smithsonian Institution, USA), for help accessing the collections. This work was supported by the Natural Science and Engineering Research Council of Canada Graduate Scholarship (NSERC CGS-D) and the Izaak Walton Killam Memorial Scholarship (Canada) to JL. Partial funding for this project was provided to JL by the Kay Ball Memorial Graduate Student Research Travel Award (Canada), the University of Alberta President’s Doctoral Prize of Distinction (Canada), the Alberta Society of Professional Biologists Graduate Scholarship (Canada), the Andrew Stewart Memorial Graduate Prize (Canada), the Devendra Jindal Graduate Scholarship (Canada), the Fondo Corrigan-ACGGP-ARES (Colombia), and additional support to JL via an NSERC Dicovery Grant RGPIN 04863 to A. Richard Palmer. JL, RWP, and AAK thank the PCP-PIRE NSF project 0966884 (OISE, EAR, DRL). CES thanks the NSF grants EF-0531670 and EAR-1223206 to Feldmann and CES. WS thanks Dione Seripieri (Museu de Zoologia, Universidade de São Paulo) for providing literature items; Antônio Carlos Sequeira Fernandes (MNRJ, Brazil) for making material from his institution available for study and for providing working space, and the FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo) for supporting studies on the systematics of decapod crustaceans (grant 2013-01201-0). RWP thanks the McGinty Endowment at the FLMNH. AAK thanks the Jon L. and Beverly A. Thompson Endowment Fund. Our most sincere thanks to Sylvain Charbonnier and Matúš Hyžný for their reviews and suggestions to improve the manuscript, and to Paula Araujo and Célio Magalhães for editorial assistance. This is Fossil Crustacea from tropical America Contribution No. 5, Panamá Decapoda Project Contribution No. 1, and University of Florida Contribution to Paleobiology No. 822.
Author contributions. JL conceived and designed the study, analyzed the data, wrote the paper, and made figs 1-4, 6-10, and Chart 1. WS made fig. 5; RWP figs 11, 12, 14-16; FJV fig. 13. All authors contributed data and edited the manuscript.
Additional information. Competing financial interests. The authors declare no competing financial interests.
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