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Nauplius

Print version ISSN 0104-6497On-line version ISSN 2358-2936

Nauplius vol.25  Cruz das Almas  2017  Epub Oct 19, 2017

http://dx.doi.org/10.1590/2358-2936e2017025 

Original Article

Checklist of fossil decapod crustaceans from tropical America. Part I: Anomura and Brachyura

Javier Luque1  2 
http://orcid.org/0000-0002-4391-5951

Carrie E. Schweitzer3 

William Santana4 
http://orcid.org/0000-0003-3086-4419

Roger W. Portell5 

Francisco J. Vega6 

Adiël A. Klompmaker7 

1 Department of Biological Sciences, University of Alberta. Edmonton, Alberta T6G 2E9, Canada.

2 Smithsonian Tropical Research Institute. Balboa-Ancón 0843-03092, Panamá, Panamá.

3 Department of Geology, Kent State University at Stark. 6000 Frank Ave. NW, North Canton, Ohio 44720, USA.

4 Universidade do Sagrado Coração - USC, Laboratório de Sistemática Zoológica. Rua Irmã Arminda, 10-50, Jd. Brazil. 17011-160 Bauru, São Paulo, Brazil.

5 Florida Museum of Natural History. 1659 Museum Road, University of Florida, Gainesville, FL 32611, USA.

6 Instituto de Geología, Universidad Nacional Autónoma de México. Coyoacán. 04510, Ciudad de México, Mexico

7 Department of Integrative Biology and Museum of Paleontology, University of California, Berkeley. 1005 Valley Life Sciences Building #3140, Berkeley, California 94720, USA.

Abstract

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.

Figure 1 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.

Chart 1 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.

Figure 2 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 DECAPODALatreille, 1802

Clade MEIURASaint Laurent, 1980

Northern South America

BOLIVIA

Infraorder ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 1892

Diogenidae incertae sedis ENT#091;Cenomanian, PotosíENT#093; (Fig. 3A-H) ENT#091;Notes 1, 2ENT#093;.

Figure 3 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 BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily ?CARPILIOIDEAOrtmann, 1893

?TumidocarcinidaeSchweitzer, 2005a

DynomenopsisSecretan, 1972

Dynomenopsis branisaiSecretan, 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)

Figure 4 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 ANOMURAMacLeay, 1838

Superfamily GALATHEOIDEASamouelle, 1819

MunidopsidaeOrtmann, 1898

BrazilomunidaMartins-Neto, 2001

Brazilomunida brasiliensis † (Beurlen, 1965, as Galatheites brasiiensis) (type) ENT#091;Albian, SergipeENT#093;

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 1892

Dardanus Paul’son, 1875

Dardanus spp. ENT#091;Paleocene, PernambucoENT#093; (inTávora et al., 2005) ENT#091;Note 1ENT#093;

Infraorder BRACHYURALatreille, 1802

Superfamily DROMIOIDEADe Haan, 1833

DynomenidaeOrtmann, 1892

MaurimiaMartins-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

AraripecarcinusMartins-Neto, 1987

Araripecarcinus ferreiraiMartins-Neto, 1987 (type) ENT#091;early Albian, CearáENT#093; (Luque, 2015b) ENT#091;Note 2ENT#093;.

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RaninoidinaeLőrenthey 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatella Smith, 1869a

Hepatella amazonicaBeurlen, 1958a ENT#091;early Miocene, ParáENT#093;.

CalappidaeDe 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).

Figure 5 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. zurcheriBouvier, 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 LEUCOSIOIDEASamouelle, 1819

Leucosiidae incertae sedis

TypilobusStoliczka, 1871

Typilobus unispinatusMartins-Neto, 2001 ENT#091;Miocene, ParáENT#093;. (as Randallia sp. in Beurlen, 1958a)

Superfamily PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 1838

Parthenope Weber, 1795

Parthenope trituberculataBeurlen, 1958a ENT#091;Miocene, ParáENT#093;.

Superfamily CANCROIDEALatreille, 1802

CancridaeLatreille, 1802

CancrinaeLatreille, 1802

CyclocancerBeurlen, 1958a

Cyclocancer tuberculatusBeurlen, 1958a (type) ENT#091;early Miocene, ParáENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

MacropipidaeStephenson and Campbell, 1960

OphthalmoplaxRathbun, 1935b

Ophthalmoplax brasiliana † (Maury, 1930, as Zanthopsis brasiliana) ENT#091; Campanian-Maastrichtian, ParaíbaENT#093; (Pralon et al. 2017) (Fig. 5E, F).

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Necronectes † A.Milne-Edwards, 1881

Necronectes tajinensisVega, 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 costataRathbun, 1919 ENT#091;Miocene, ParáENT#093; (reported in Beurlen, 1958a, and Távora et al., 2002).

PodophthalminaeDana, 1851d

Euphylax Stimpson, 1862

Euphyplax septendentatusBeurlen, 1958a ENT#091; early Miocene, ParáENT#093;.

PortuninaeRafinesque, 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 paraensisBeurlen, 1958a ENT#091;early Miocene, ParáENT#093; (Fig. 5C, D).

Callinectes reticulatusRathbun, 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 haitensisRathbun, 1923a ENT#091;Miocene, ParáENT#093; (sensuTávora et al., 2002).

Portunus pirabaensisMartins-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 RETROPLUMOIDEAGill, 1894

RetroplumidaeGill, 1894

?ArchaeopusRathbun, 1908

?Archaeopus rathbunaeBeurlen, 1965 ENT#091;Albian, SergipeENT#093; ENT#091;Note 7ENT#093;.

CostacoplumaCollins and Morris, 1975

Costacopluma nordestinaFeldmann 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 CARPILIOIDEAOrtmann, 1893

Palaeoxanthopsidae †Schweitzer, 2003

PalaeoxanthopsisBeurlen, 1958b

Palaeoxanthopsis cretacea † (Rathbun, 1902) (type) ENT#091;Maastrichtian, ParaibaENT#093;.

Tumidocarcinidae †Schweitzer, 2005a

ParatumidocarcinusMartins-Neto, 2001

Paratumidocarcinus marajoarusMartins-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 XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 1893

Panopeus H.Milne Edwards, 1834

Panopeus capanemaensisMartins-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 THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 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;.

Figure 6 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. 

SesarmidaeDana, 1851c

Sesarma Say, 1817

Sesarma paraensisBeurlen, 1958a ENT#091;Miocene, ParáENT#093;.

Superfamily OCYPODOIDEARafinesque, 1815

OcypodidaeRafinesque, 1815

Uca Leach, 1814

Uca antiquaBrito, 1972 (junior synonym U. inaciobritoiMartins-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 ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

?Paguroidea incertae sedis ENT#091;Maastrichtian, AlgarroboENT#093; (Schweitzer et al., 2006a).

Infraorder BRACHYURALatreille, 1802

Superfamily HOMOLOIDEADe Haan, 1833

HomolidaeDe Haan, 1839

HomolopsisBell, 1863

Homolopsis chilensisFörster and Stinnesbeck, 1987 ENT#091;Maastrichtian, near ConcepciónENT#093;.

Superfamily RANINOIDEADe Haan, 1839

LyreididaeGuinot, 1993

LyreidinaeGuinot, 1993

Lyreidus De Haan, 1841

Lyreidus lebuensisFeldmann, 1992 ENT#091;Eocene, near LebuENT#093; (as Lyreidus sp. inFeldmann and Chirino-Galvez, 1991).

RaninidaeDe Haan, 1839

RaninoidinaeLőrenthey in Lőrenthey and Beurlen, 1929

Raninoides H.Milne Edwards, 1837

Raninoides araucana † (Philippi, 1887, as Symnista araucana) ENT#091;Eocene, near LebuENT#093;.

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus spinimarginatusFeldmann, Schweitzer and Encinas, 2005 ENT#091;Miocene, Cardenal CaroENT#093;.

CalappidaeDe Haan, 1833

Calappilia † A.Milne-Edwards, 1873

?Calappilia chilensisFeldmann, Schweitzer and Encinas, 2005 ENT#091;Miocene, Cardenal CaroENT#093;.

Superfamily LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 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;.

Figure 7 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 CANCROIDEALatreille, 1802

AtelecyclidaeOrtmann, 1893

Trichopeltarion A.Milne-Edwards, 1880b

Trichopeltarion frassinettiFeldmann, Schweitzer and Encinas, 2010 ENT#091;late Pliocene, Guafo IslandENT#093;.

Trichopeltarion levisCasadío et al., 2004 ENT#091;Miocene, Cardenal CaroENT#093; (Feldmann et al., 2005; 2010).

Superfamily CHEIRAGONOIDEAOrtmann, 1893

CheriagonidaeOrtmann, 1893

PirulellaFeldmann, Schweitzer and Encinas, 2010

Pirulella antipodeaFeldmann, Schweitzer and Encinas, 2010 ENT#091;early Pliocene, Chiloé IslandENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

GeryonidaeColosi, 1923

ArchaeogeryonColosi, 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 quadrataFeldmann, 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;.

MacropipidaeStephenson and Campbell, 1960

MinohellenusKarasawa, 1990

Minohellenus araucanus † (Philippi, 1887) ENT#091;Miocene, locality indet.ENT#093;.

PortunidaeRafinesque, 1815

PheophthalmusFeldmann, Schweitzer and Encinas, 2010

Pheophthalmus mochaensisFeldmann, Schweitzer and Encinas, 2010 ENT#091;Miocene, Mocha IslandENT#093;.

Superfamily GONEPLACOIDEAMacLeay, 1838

ChasmocarcinidaeSerène, 1964

ChasmocarcininaeSerène, 1964

Chasmocarcinus Rathbun, 1898

Chasmocarcinus chiloeensisFeldmann, Schweitzer and Encinas, 2010 ENT#091;Miocene, Chiloé IslandENT#093;.

Superfamily PILUMNOIDEASamouelle, 1819

PilumnidaeSamouelle, 1819

PilumninaeSamouelle, 1819

Pilumnus Leach, 1816

Pilumnus cucaoensisFeldmann, Schweitzer and Encinas, 2005 ENT#091;Miocene, Chiloé Island and Cardenal CaroENT#093; (Feldmann et al., 2010).

Subsection THORACOTREMATAGuinot, 1977

Superfamily PINNOTHEROIDEADe Haan, 1833

PinnotheridaeDe Haan, 1833

PinnothereliinaeAlcock, 1900a

Pinnixa White, 1846

Pinnixa navidadensisFeldmann, 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 ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 1892

?Paguristes Dana, 1851c

?Paguristes sp. ENT#091;late Valanginian, SantanderENT#093; (Fig. 8A) ENT#091;Note 1ENT#093;.

Figure 8 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 BRACHYURALatreille, 1802

Superfamily DROMIOIDEADe 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 HOMOLOIDEADe Haan, 1833

Mithracitidae †Števčić, 2005

MithracitesGould, 1859

Mithracites takedaiVan Bakel, Guinot, Jagt and Fraaije, 2012b ENT#091;late Aptian, SantanderENT#093;

Superfamily NECROCARCINOIDEA †Förster, 1968

Cenomanocarcinidae †Guinot, Vega and Van Bakel, 2008

CenomanocarcinusVan Straelen, 1936

Cenomanocarcinus vanstraeleniStenzel, 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

BellcarcinusLuque, 2014

Bellcarcinus aptiensisLuque, 2014 (type) ENT#091;late Aptian, Santander and BoyacáENT#093; ENT#091;Note 3ENT#093;.

ColombicarcinusKarasawa, Schweitzer, Feldmann and Luque, 2014.

Colombicarcinus laevisKarasawa, Schweitzer, Feldmann and Luque, 2014 (type) ENT#091;late Aptian, Santander and BoyacáENT#093; ENT#091;Note 3ENT#093;.

PlanocarcinusLuque, 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 johnjagtiBermúdez, 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

JoeraninaVan 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 colombianaBermúdez, Crúz and Vega in Bermúdez et al., 2013 ENT#091;early-middle Albian, BoyacáENT#093;.

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RaninoidinaeLőrenthey in Lőrenthey and Beurlen, 1929

QuasilaeviraninaTucker, 1998

Quasilaeviranina sp. † ENT#091;early-middle Santonian to Maastrichtian of Boyacá, Tolima, and CundinamarcaENT#093; (Fig. 8C) ENT#091;Note 4ENT#093;.

Section EUBRACHYURASaint 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 HETEROTREMATAGuinot, 1977

Superfamily DORIPPOIDEAMacLeay, 1838

Telamonocarcinidae †Larghi, 2004

TelamonocarcinusLarghi, 2004

Telamonocarcinus antiquusLuque, 2015a ENT#091;early Albian, SantanderENT#093; (Fig. 8D) ENT#091;Note 7ENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

MacropipidaeStephenson and Campbell, 1960

OphthalmoplaxRathbun, 1935b

Ophthalmoplax andinaGuzmán, 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;.

PortunidaeRafinesque, 1815

PortuninaeRafinesque, 1815

Callinectes Stimpson, 1862

Callinectes reticulatusRathbun, 1918 (claw) ENT#091;Oligocene, AntioquiaENT#093;.

Portunus Weber, 1795

Portunus oblongusRathbun, 1920b ENT#091;Miocene, La GuajiraENT#093; (Fig. 8F, G) ENT#091;Note 9ENT#093;.

Superfamily RETROPLUMOIDEAGill, 1894

RetroplumidaeGill, 1894

CostacoplumaCollins 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

DilocarcininaePretzmann, 1978

Sylviocarcinus H.Milne Edwards, 1853

Sylviocarcinus piriformis (Pretzmann, 1968) ENT#091;early Miocene, TolimaENT#093; (Fig. 4B) ENT#091;Note 11ENT#093;.

Superfamily CARPILIOIDEAOrtmann, 1893

Palaeoxanthopsidae †Schweitzer, 2003

PalaeoxanthopsisBeurlen, 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 BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Necronectes † A.Milne-Edwards, 1881

Necronectes proavitus † (Rathbun, 1918) ENT#091;early Miocene, Cuenca BasinENT#093; (Morris, 1973) ENT#091;Note 1ENT#093;.

PortuninaeRafinesque, 1815

Portunus Weber, 1795

Portunus oblongusRathbun, 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 BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

EpialtidaeMacLeay, 1838

EpialtinaeMacLeay, 1838

EoinachoidesVan Straelen, 1933b

Eoinachoides latispinosaCarriol, Muizon and Secretán, 1987 ENT#091;late Miocene, PiscoENT#093; (Fig. 9C).

Figure 9 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. 

PisinaeDana, 1851b

Libinia Leach, 1815

Libinia peruviensisCarriol, Muizon and Secretán, 1987 ENT#091;early Pliocene, PiscoENT#093;.

Superfamily CANCROIDEALatreille, 1802

CancridaeLatreille, 1802

CancrinaeLatreille, 1802

Cancer Linnaeus, 1758

Cancer borealisStimpson, 1862 ENT#091;Miocene-early Pliocene, Pisco and ArequipaENT#093; (Fig. 9A, B).

Superfamily PORTUNOIDEARafinesque, 1815

GeryonidaeColosi, 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;.

PortunidaeRafinesque, 1815

PodophthalminaeDana, 1851d

Euphylax Stimpson, 1862

Euphylax sp. cf. E. callinectiasRathbun, 1918 ENT#091;Eocene, PiscoENT#093; (Fig. 9F-H) ENT#091;Notes 2, 3ENT#093;.

PortuninaeRafinesque, 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 ?reticulatusRathbun, 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 CARPILIOIDEAOrtmann, 1893

Zanthopsidae †Vía Boada, 1959

ZanthopsisM'Coy, 1849

Zanthopsis erransWoods, 1922 ENT#091;Eocene, PiuraENT#093;.

Superfamily HEXAPODOIDEAMiers, 1886

HexapodidaeMiers, 1886

PalaeopinnixaVía 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 ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 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 BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RaninoidinaeLőrenthey in Lőrenthey and Beurlen, 1929

Raninoides H.Milne Edwards, 1837

Raninoides rathbunaeVan 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

EpialtidaeMacLeay, 1838

EpialtinaeMacLeay, 1838

EoinachoidesVan Straelen, 1933b

Eoinachoides senniVan 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).

MithracidaeMacLeay, 1838

Mithrax Desmarest, 1823

Mithrax sp. ENT#091;Plio-Pleistocene, Falcón and SucreENT#093; (Aguilera et al., 2010).

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

EriosachilaBlow 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 amazonicaBeurlen, 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;.

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa laraensisVan 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 LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 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 PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 1838

Parthenope Weber, 1795

Parthenope venezuelensisVan 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 PORTUNOIDEARafinesque, 1815

MacropipidaeStephenson and Campbell, 1960

OphthalmoplaxRathbun, 1935b

Ophthalmoplax brasiliana † (Maury, 1930 ENT#091;junior synonym O. triambonatusFeldmann and Villamil, 2002ENT#093;) ENT#091;Maastrichtian, TáchiraENT#093; (Aguilera et al., 2010).

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 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;.

PodophthalminaeDana, 1851d

Euphylax Stimpson, 1862

Euphylax sp. ENT#091;Miocene to Pliocene, Falcón and ZuliaENT#093;.

Paraeuphylax † Varela andSchweitzer, 2011

Paraeuphyplax cubaensis † Varela and Schweitzer, 2011 ENT#091;early Miocene, ZuliaENT#093; (Fig. 10I) ENT#091;Note 3ENT#093;.

Figure 10 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. 

?SaratunusCollins, Lee and Noad, 2003

?Saratunus sp. † ENT#091;early Miocene, FalcónENT#093; (Aguilera et al., 2010) ENT#091;Note 2ENT#093;.

PortuninaeRafinesque, 1815

Callinectes Stimpson, 1862

Callinectes reticulatusRathbun, 1918 (claws only) ENT#091;early Miocene, ZuliaENT#093; (Fig. 10E, F).

Callinectes sp. cf. C. declivisRathbun, 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 oblongusRathbun, 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 RETROPLUMOIDEAGill, 1894

RetroplumidaeGill, 1894

CostacoplumaCollins and Morris, 1975

Costacopluma bifidaCollins, 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 GONEPLACOIDEAMacLeay, 1838

ChasmocarcinidaeSerène, 1964

ChasmocarcininaeSerène, 1964

FalconoplaxVan Straelen, 1933b

Falconoplax kugleriVan Straelen, 1933b (type) ENT#091;middle-late Eocene to early Miocene, Lara and FalcónENT#093; (Feldmann and Schweitzer, 2004; Aguilera et al., 2010).

Superfamily HEXAPODOIDEAMiers, 1886

HexapodidaeMiers, 1886

PalaeopinnixaVía Boada, 1966

Palaeopinnixa perornataCollins 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 XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 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 BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily PORTUNOIDEARafinesque, 1815

Carcineretidae †Beurlen, 1930

CarcineretesWithers, 1922

Carcineretes planetariusVega, Feldmann, Ocampo and Pope, 1997 ENT#091;Maastrichtian, Albion IslandENT#093;.

COSTA RICA

Infraorder ANOMURAMacLeay, 1838

Superfamily GALATHEOIDEASamouelle, 1819

PorcellanidaeHaworth, 1825

Pachycheles Stimpson, 1858

Pachycheles latusRathbun, 1918 ENT#091;Pliocene, Port LimónENT#093; (Fig.11A).

Figure 11 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 avitusRathbun, 1918 ENT#091;Pliocene, Port LimónENT#093; (Fig.11B).

Infraorder BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RaninoidinaeLőrenthey in Lőrenthey and Beurlen, 1929

Raninoides H.Milne Edwards, 1837

Raninoides sp. cf. R. benedictiRathbun, 1935a ENT#091;?early Pleistocene, Burica PeninsulaENT#093;.

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

MithracidaeMacLeay, 1838

Thoe Bell, 1836

Thoe asperoidesCollins and Todd in Todd and Collins, 2005 ENT#091;late Pliocene to early Pleistocene, LimónENT#093;.

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus lineatinusCollins and Todd in Todd and Collins, 2005 ENT#091;late Pliocene to early Pleistocene, Puntarenas).

Hepatus biformisCollins in Todd and Collins, 2005 ENT#091;early Pliocene, PuntarenasENT#093;.

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa costaricanaRathbun, 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 LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 1871

Leucosilia Bell, 1855

Leucosilia bananensisRathbun, 1918 ENT#091;Pliocene, Limón)ENT#093; (Todd and Collins, 2005).

Speleophorus A.Milne-Edwards, 1865

Speleophorus ?subcircularisCollins 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 PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

Platylambrus Stimpson, 1871

Platylambrus spinulatusCollins and Todd in Todd and Collins, 2005 ENT#091;early Pleistocene, LimónENT#093;

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

PodophthalminaeDana, 1851d

Euphylax Stimpson, 1862

Euphyplax callinectiasRathbun, 1918 ENT#091;early Pliocene (not Miocene), LimónENT#093; (Todd and Collins, 2005).

Euphyplax fortisRathbun, 1918 ENT#091;early Pliocene (not Miocene), LimónENT#093; (Todd and Collins, 2005).

Euphyplax maculatusCollins and Todd in Todd and Collins, 2005 (claw fragments) ENT#091;early Pleistocene, PuntarenasENT#093;.

SandomingiaRathbun, 1919

Sandomingia yaquiensisRathbun, 1919 ENT#091;late Pliocene, LimónENT#093;.

PortuninaeRafinesque, 1815

Callinectes Stimpson, 1862

Callinectes declivisRathbun, 1918 (fingers only) ENT#091;early Pliocene to early Pleistocene, LimónENT#093; (Todd and Collins, 2005).

Portunus Weber, 1795

Portunus gabbiRathbun, 1919 ENT#091;Pliocene, LimónENT#093;.

Superfamily XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 1893

Lophopanopeus Rathbun, 1898

Lophopanopeus maculoidesCollins and Todd in Todd and Collins, 2005 ENT#091;late Pliocene, LimónENT#093;.

XanthidaeMacLeay, 1838

ActaeinaeAlcock, 1898

Heteractaea Lockington, 1877

Heteractaea lunata (H. Milne Edwards and Lucas, 1843) ENT#091;late Pliocene to early Pleistocene, LimónENT#093;.

Subsection THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 1838

Cardisoma Latreille, 1828

Cardisoma guanhumiLatreille, 1828 ENT#091;late Pliocene to early Pleistocene, LimónENT#093;.

HONDURAS

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection THORACOTREMATAGuinot, 1977

Superfamily OCYPODOIDEARafinesque, 1815

OcypodidaeRafinesque, 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 ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 1892

Dardanus Paul’son, 1875

Dardanus biordinesCollins and Todd in Todd and Collins, 2005 ENT#091;early Pliocene, Bocas del ToroENT#093;.

Petrochirus Stimpson, 1858

Petrochirus bouvieriRathbun, 1918 ENT#091;late Miocene to late Pliocene, Bocas del Toro, Colón and DariénENT#093;.

Infraorder BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RanininaeDe Haan, 1839

?Ranina Lamarck, 1801

?Ranina sp. ENT#091;Miocene, ChiriquíENT#093;.

RaninoidinaeLőrenthey 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily DORIPPOIDEAMacLeay, 1838

?Goniochelidae †Schweitzer and Feldmann, 2011

?GoniocheleBell, 1858

?Goniochele armataRathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093; (Fig. 11E) ENT#091;Note 1ENT#093;.

Superfamily MAJOIDEASamouelle, 1819

MithracidaeMacLeay, 1838

Thoe Bell, 1836

Thoe asperoidesCollins and Todd in Todd and Collins, 2005 ENT#091;late Miocene, Bocas del ToroENT#093;.

OregoniidaeGarth, 1958

Hyas Leach, 1814 ENT#091;inLeach, 1813-1815ENT#093;

Hyas sp. ENT#091;late Miocene, DariénENT#093; (Todd and Collins, 2005).

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

EriosachilaBlow and Manning, 1996

Eriosachila terryi † (Rathbun, 1937, as Zanthopsis terryi) ENT#091;late Eocene, PanamaENT#093;.

Hepatus Latreille, 1802

Hepatus biformisCollins 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 lineatinusCollins 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).

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa flammea (Herbst, 1794) ENT#091;late Miocene to Pleistocene, ColónENT#093;.

Calappa zurcheriBouvier, 1899 ENT#091;Miocene, unknown provenanceENT#093; (Fig. 12M) ENT#091;Note 2ENT#093;.

Figure 12 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).

CalappellaRathbun, 1918

Calappella quadrispinaRathbun, 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 macdonaldiRathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.

Mursia obscuraRathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.

MursiliaRathbun, 1918

Mursilia ecristataRathbun, 1918 ENT#091;early to middle Miocene, ColónENT#093;.

Superfamily LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 1871

Iliacantha Stimpson, 1871

Iliacantha panamicaCollins 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 bananensisRathbun, 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 enigmaticaCollins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.

Persephona manningiCollins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.

Speloeophorus A.Milne-Edwards, 1865

Speloeophorus subcircularisCollins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.

Superfamily PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 1838

Parthenope Weber, 1795

Parthenope panamensisRathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.

Parthenope pleistocenicaRathbun, 1918 ENT#091;Pleistocene, ColónENT#093;.

Platylambrus Stimpson, 1871

Platylambrus spinulatusCollins and Todd in Todd and Collins, 2005 ENT#091;early late Pliocene, Bocas del ToroENT#093;.

Superfamily CANCROIDEALatreille, 1802

CancridaeLatreille, 1802

CancrinaeLatreille, 1802

Cancer Latreille, 1802

Cancer santosi † (Rathbun, 1937, as Lobocarcinus santosi) ENT#091;late Eocene, Los SantosENT#093;.

Superfamily CHEIRAGONOIDEAOrtmann, 1893

CheriagonidaeOrtmann, 1893

MontezumellaRathbun, 1930

Montezumella casayetensisRathbun, 1937 ENT#091;late Oligocene or early Miocene, Bahía de PanamáENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Necronectes † A.Milne-Edwards, 1881

Necronectes proavitus † (Rathbun, 1918, as Gatunia proavita) ENT#091;early Miocene, ColónENT#093; (Fig. 12N).

PodophthalminaeDana, 1851d

Euphylax Stimpson, 1862

Euphylax callinectiasRathbun, 1918 ENT#091;early Miocene (not Oligocene), Panama CanalENT#093;.

Euphylax maculatusCollins 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).

SandomingiaRathbun, 1919

Sandomingia yaquiensisRathbun, 1919 ENT#091;early to late Pliocene, Bocas del ToroENT#093;.

PortuninaeRafinesque, 1815

Callinectes Stimpson, 1862

Callinectes declivisRathbun, 1918 ENT#091;late Eocene to early Miocene, Panama CanalENT#093;.

Callinectes reticulatusRathbun, 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 gabbiRathbun, 1919 ENT#091;middle-late Miocene, DariénENT#093;.

Portunus spp. cf. P. tenuisRathbun, 1919 ENT#091;Pleistocene, ChiriquíENT#093; ENT#091;Note 3ENT#093;.

Superfamily ERIPHIOIDEAMacLeay, 1838

EriphiidaeMacLeay, 1838

Eriphia Latreille, 1817

Eriphia sp. aff. E. squamataStimpson, 1859 ENT#091;Quaternary, Bahía de PanamáENT#093; (Luque et al., in press).

PlatyxanthidaeGuinot, 1977

Platyxanthus A.Milne-Edwards, 1863

Platyxanthus sp. ENT#091;late Miocene, Bocas del ToroENT#093; (Todd and Collins, 2005).

Superfamily GONEPLACOIDEAMacLeay, 1838

ChasmocarcinidaeSerène, 1964

ChasmocarcininaeSerène, 1964

FalconoplaxVan Straelen, 1933b

Falconoplax kugleriVan Straelen, 1933b ENT#091;early Miocene, Panama CanalENT#093; (Fig. 11J-Q) ENT#091;Note 3ENT#093;.

EuryplacidaeStimpson, 1871

Euryplax Stimpson, 1859

? Euryplax culebrensisRathbun, 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 HEXAPODOIDEAMiers, 1886

HexapodidaeMiers, 1886

PalaeopinnixaVía 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 PILUMNOIDEASamouelle, 1819

PilumnidaeSamouelle, 1819

PilumninaeSamouelle, 1819

Pilumnus Leach, 1816

Pilumnus sp. ENT#091;Pliocene - early Pleistocene, Bocas del Toro and ChiriquíENT#093; (Todd and Collins, 2005).

Superfamily XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 1893

Eurytium Stimpson, 1859

Eurytium crenulatumRathbun, 1918 ENT#091;Pleistocene, ColónENT#093; ENT#091;Note 7ENT#093;.

Panopeus H.Milne Edwards, 1834

Panopeus antepurpureusRathbun, 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).

XanthidaeMacLeay, 1838

ActaeinaeAlcock, 1898

Heteractaea Lockington, 1877

Heteractaea lunata (H. Milne Edwards and Lucas, 1843) ENT#091;late Pliocene, Bocas del ToroENT#093;.

XanthinaeMacLeay, 1838

Micropanope Stimpson, 1871

Micropanope sp. ENT#091;early late Pliocene, Bocas del ToroENT#093; (Todd and Collins, 2005).

Subsection THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 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;.

GrapsidaeMacLeay, 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 OCYPODOIDEARafinesque, 1815

OcypodidaeRafinesque, 1815

OcypodinaeRafinesque, 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 PINNOTHEROIDEADe Haan, 1833

PinnotheridaeDe Haan, 1833

PinnothereliinaeAlcock, 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 ANOMURAMacLeay, 1838

Superfamily AEGLOIDEADana, 1852

AeglidaeDana, 1852

ProtaeglaFeldmann, Vega, Applegate and Bishop, 1998b

Protaegla minisculaFeldmann, Vega, Applegate and Bishop, 1998b (type) ENT#091;Albian, PueblaENT#093; (Vega et al., 2005) (Fig. 13A).

Figure 13 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 GALATHEOIDEASamouelle, 1819

GalatheidaeSamouelle, 1819

Galatheidae indet. ENT#091;middle Eocene, Baja California SurENT#093; (Schweitzer et al., 2006b).

Superfamily PAGUROIDEALatreille, 1802

Paguroidea spp. indet. ENT#091;middle Eocene, Baja California SurENT#093; (Schweitzer et al., 2005).

DiogenidaeOrtmann, 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).

PaguridaeLatreille, 1802

Pagurus Fabricius, 1775

Pagurus sp. ENT#091;Albian, PueblaENT#093; (Vega et al., 2005).

PalaeopagurusVan Straelen, 1925

Palaeopagurus sp. cf. P. pilsbyiRoberts, 1962 ENT#091;early Maastrichtian, Nuevo LeónENT#093; (Vega et al., 1995b).

Infraorder BRACHYURALatreille, 1802

Superfamily DROMIOIDEADe Haan, 1833

DynomenidaeOrtmann, 1892

GraptocarcinusRoemer, 1887

Graptocarcinus muiriStenzel, 1944b ENT#091;Albian, San Luis PotosíENT#093;.

Xandarocarcinidae †Karasawa Schweitzer and Feldmann, 2011

XandarocarcinusKarasawa, 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

PrehepatusRathbun, 1935b

Prehepatus harrisiBishop, 1985 ENT#091;early Maastrichtian, Nuevo LeónENT#093; (Vega et al., 1995b).

Prehepatus mexicanusSchweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.

Superfamily HOMOLOIDEADe Haan, 1833

HomolidaeDe Haan, 1839

Homola Leach, 1816 ENT#091;imprint 1815ENT#093;

Homola bajaensisSchweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.

ZygastrocarcinusBishop, 1983a

Zygastrocarcinus carolinasensisKlompmaker, 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

CenomanocarcinusVan Straelen, 1936

Cenomanocarcinus vanstraeleniStenzel, 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

FerroraninaVan Bakel, Guinot, Artal, Fraaije and Jagt, 2012a

Ferroranina sp. cf. F. dichrous † (Stenzel, 1945) ENT#091;Turonian, CoahuilaENT#093; (Vega et al., 2007b).

Superfamily RANINOIDEADe Haan, 1839

LyreididaeGuinot, 1993

Macroacaeninae †Karasawa, Schweitzer, Feldmann and Luque, 2014

MacroacaenaTucker, 1998

Macroacaena venturaiVega, Nyborg, Fraaye and Espinosa, 2007a ENT#091;middle Paleocene (Selandian), Chiapas and CoahuilaENT#093;.

Marylyreidinae †Van Bakel, Guinot, Artal, Fraaije and Jagt, 2012a

BournelyreidusVan Bakel, Guinot, Artal, Fraaije and Jagt, 2012a

Bournelyreidus oaheensis † (Bishop, 1978, as Raninella oaheensis) ENT#091;early Maastrichtian, CoahuilaENT#093; (Vega et al., 2016).

RaninidaeDe Haan, 1839

Raninidae spp. indet. ENT#091;Turonian, CoahuilaENT#093; (Vega et al., 2007b).

CyrtorhininaeGuinot, 1993

ClaudioraninaKarasawa, Schweitzer, Feldmann and Luque, 2014

Claudioranina latacanthaMartínez, Aguillón, Luque and Vega, 2017 ENT#091;middle Paleocene (Selandian), CoahuilaENT#093;.

NotopodinaeSerène and Umali, 1972

Notopus De Haan, 1841

Notopus minutusVega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a ENT#091;Ypresian, ChiapasENT#093;.

RanininaeDe Haan, 1839

LophoraninaFabiani, 1910

Lophoranina bishopiSquires and Demetrion, 1992 ENT#091;Ypresian and Eocene indet., Baja California, SurENT#093; (Schweitzer et al., 2002; 2007b).

Lophoranina cristaspinaVega, 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 berglundiSquires 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)

VegaraninaVan 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;.

RaninoidinaeLőrenthey in Lőrenthey and Beurlen, 1929

Notopoides Henderson, 1888

Notopoides exiguusBeschin, 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).

QuasilaeviraninaTucker, 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 acanthocolusSchweitzer, Feldmann, Gonzalez-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.

Raninoides mexicanusRathbun, 1930 ENT#091;Miocene, VeracruzENT#093;.

Raninoides proracanthusSchweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.

Raninoides treldenaesensisCollins and Jakobsen, 2003 ENT#091;Ypresian, ChiapasENT#093; (Vega et al., 2008).

Section DAKOTICANCROIDA †Rathbun, 1917

Superfamily DAKOTICANCROIDEARathbun, 1917

Dakoticancridae †Rathbun, 1917

DakoticancerRathbun, 1917

Dakoticancer australisRathbun, 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

?SodakusBishop, 1978

Sodakus mexicanusVega, Feldmann and Villalobos-Hiriart, 1995b ENT#091;early Maastrichtian, Nuevo León and CoahuilaENT#093; (Vega et al., 2016) (Fig. 13D).

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily ?DORIPPOIDEAMacLeay, 1838

Tepexicarcinidae †Luque, 2015a

TepexicarcinusFeldmann, Vega, Applegate and Bishop, 1998b

Tepexicarcinus tlayuaensisFeldmann, Vega, Applegate and Bishop, 1998b ENT#091;Albian, PueblaENT#093; (Vega et al., 2005) (Fig. 13E).

Superfamily MAJOIDEASamouelle, 1819

OregoniidaeGarth, 1958

Oregonia Dana, 1851a

MajidaeSamouelle, 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 CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

EriosachilaBlow and Manning, 1996

Eriosachila bajaensisSchweitzer, 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 amazonicaBeurlen, 1958a ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa flammea (Herbst, 1794) ENT#091;Oligocene, Baja California SurENT#093; (Rathbun, 1930).

Calappa zurcheriBouvier, 1899 ENT#091;Oligocene, Veracruz; early Miocene, ChiapasENT#093; (Rathbun, 1930; Vega et al., 2009).

Calappilia † A.Milne-Edwards, 1873

Calappilia hondoensisRathbun, 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 LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 1871

Iliacantha Stimpson, 1871

Iliacantha panamanicaCollins and Todd in Todd and Collins, 2005 ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).

Superfamily PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

DaldorfiinaeNg and Rodríguez, 1986

Daldorfia Rathbun, 1904

Daldorfia salinaSchweitzer, Feldmann, González-Barba and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.

Superfamily CANCROIDEALatreille, 1802

AtelecyclidaeOrtmann, 1893

LevicyclusSchweitzer, Feldmann, Gonzáles-Barba and Vega, 2002

Levicyclus tepetateSchweitzer, Feldmann, González-Barba, and Vega, 2002 ENT#091;Eocene, Baja California SurENT#093;.

CancridaeLatreille, 1802

CancrinaeLatreille, 1802

Anatolikos Schweitzer and Feldmann, 2000

Anatolikos undecimspinosusSchweitzer, 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 CHEIRAGONOIDEAOrtmann, 1893

CheriagonidaeOrtmann, 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).

MontezumellaRathbun, 1930

Montezumella tubulataRathbun, 1930 ENT#091;Eocene: Priabonian, Baja California SurENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

Carcineretidae †Beurlen, 1930

CarcineretesWithers, 1922

Carcineretes planetariusVega, Feldmann, Ocampo and Pope, 1997 ENT#091;early Maastrichtian, ChiapasENT#093; (Vega et al., 2001b) (Fig. 13I).

CarcinidaeMacLeay, 1838

?Xaiva MacLeay, 1838

?Xaiva sp. † ENT#091;middle Eocene, ChiapasENT#093; (Vega et al., 2008).

Icriocarcinidae †Števčić, 2005

IcriocarcinusBishop, 1988

Icriocarcinus xestosBishop, 1988 ENT#091;Maastrichtian, Baja California SurENT#093; (Schweitzer et al., 2002; Phillips et al., 2014) (Fig. 13H).

BranchiocarcinusVega, Feldmann and Sour-Tovar, 1995a

Branchiocarcinus cornatusFeldmann 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

LongusorbisRichards, 1975

Longusorbis eutychiusSchweitzer, Feldmann and Karasawa, 2007c ENT#091;middle Eocene (Bartonian), Baja California SurENT#093;.

Longusorbis quadratusFraaije, Vega, Van Bakel and Garibay-Romero, 2006 ENT#091;Cenomanian-Coniacian, GuerreroENT#093;.

MacropipidaeStephenson and Campbell, 1960

MaeandricampusSchweitzer and Feldmann, 2002

Maeandricampus americanus † (Rathbun, 1930, as Xanthosia americana) ENT#091;Eocene, Baja California SurENT#093;.

OphthalmoplaxRathbun, 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).

PortunidaeRafinesque, 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).

NecronectinaeGlaessner, 1928

Necronectes † A.Milne-Edwards, 1881

Necronectes nodosusSchweitzer, Feldmann, González-Barba and Vega, 2002 ENT#091;Oligocene, Baja California SurENT#093;.

Necronectes tajinensisVega et al., 1999 ENT#091;middle Miocene, VeracruzENT#093;.

Necronectes sp. † ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).

PortuninaeRafinesque, 1815

Portunus Weber, 1795

Portunus atecuicitlisVega et al., 1999 ENT#091;early to middle Miocene, Chiapas and VeracruzENT#093; (Vega et al., 2009).

Scylla De Haan, 1833

Scylla costataRathbun, 1919 ENT#091;age and locality unknownENT#093; (Rathbun, 1930).

PodophthalminaeDana, 1851d

Podophthalmus Lamarck, 1801

?Podophthalmus sp. † ENT#091;late Cretaceous, TamaulipasENT#093; (Rathbun, 1930).

Superfamily RETROPLUMOIDEAGill, 1894

RetroplumidaeGill, 1894

ArchaeopusRathbun, 1908

Archaeopus mexicanus † Schweitzer, Feldmann, González-Barba and Vega, 2002 ENT#091;Campanian-Maastrichtian, Baja California SurENT#093;.

CostacoplumaCollins and Morris, 1975

Costacopluma bishopiVega and Feldmann, 1992 ENT#091;Coniacian, GuerreroENT#093; (Fraaije et al., 2006; Martínez-Díaz et al., 2016; Vega et al., 2016).

Costacopluma grayiFeldmann and Portell, 2007 ENT#091;late Maastrichtian, CoahuilaENT#093; (Martínez-Díaz et al., 2016; Vega et al., 2016).

Costacopluma mexicanaVega 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 CARPILIOIDEAOrtmann, 1893

Palaeoxanthopsidae †Schweitzer, 2003

PalaeoxanthopsisBeurlen, 1958b

Palaeoxanthopsis meyapaquensis † (Vega, Feldmann, García-Barrera, Filkorn, Pimentel and Avendaño, 2001b, as Paraxanthopsis meyapaquensis) ENT#091;early Maastrichtian, ChiapasENT#093;.

ParaverrucoidesSchweitzer, 2003

Paraverrucoides alabamensis † (Rathbun, 1935b, as Xanthilites alabamensis) ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Vega et al., 2007a; Martínez-Díaz et al., 2017).

VerrucoidesVega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a

Verrucoides stenohedraVega, 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 mexicanusRathbun, 1930 ENT#091;Eocene, Baja California SurENT#093; (Schweitzer et al., 2002; 2006b; 2007a).

Zanthopsidae †Vía Boada, 1959

NeozanthopsisSchweitzer, 2003

Neozanthopsis americanus † (Rathbun, 1928, as Harpactocarcinus americanus) ENT#091;?middle Eocene, Baja California SurENT#093; (Vega et al., 2006b).

Superfamily GONEPLACOIDEAMacLeay, 1838

EuryplacidaeStimpson, 1871

OrbitoplaxTucker and Feldmann, 1990

Orbitoplax nandachare † (Vega et al., 2001a, as Stoaplax nandachare) ENT#091;Ypresian, ChiapasENT#093; (Vega et al., 2008).

GoneplacidaeMacLeay, 1838

Amydrocarcinus † Schweitzer,Feldmann, González-Barba and Vega, 2002

Amydrocarcinus danteiSchweitzer, Feldmann, González-Barba and Vega, 2002 ENT#091;middle Eocene (Bartonian), Baja California SurENT#093; (Schweitzer et al., 2007a).

LitocheiridaeŠtevčić, 2005

ParacorallicarcinusTessier, Beschin, Bussulini and De Angeli, 1999

Paracorallicarcinus tricarinatusSchweitzer, Feldmann, González-Barba, and Ćosović, 2006b ENT#091;middle Eocene, Baja California SurENT#093;.

MathildellidaeKarasawa and Kato, 2003

Mathildellidae indet. ENT#091;late Campanian, CoahuilaENT#093; (Vega et al., 2016).

TehuacanaStenzel, 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 schweitzeraeVega, Nyborg, Coutiño, and Hernández-Monzón, 2008 ENT#091;Eocene (Lutetian), ChiapasENT#093;.

Superfamily HEXAPODOIDEAMiers, 1886

HexapodidaeMiers, 1886

PalaeopinnixaVía Boada, 1966

Palaeopinnixa perornataCollins and Morris, 1976 ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).

Superfamily PILUMNOIDEASamouelle, 1819

PilumnidaeSamouelle, 1819

EumedoninaeDana, 1853

SanteellaBlow and Manning, 1996

Santeella lillyaeBlow and Manning, 1996 ENT#091;early Eocene (Ypresian), ChiapasENT#093; (Vega et al., 2008).

Superfamily TRAPEZIOIDEAMiers, 1886

TrapeziidaeMiers, 1886

ArchaeotetraSchweitzer, 2005b

Archaeotetra inornataSchweitzer, 2005b ENT#091;middle Eocene, Baja California SurENT#093;.

Superfamily XANTHOIDEAMacLeay, 1838

Xanthoidea indet. ENT#091;middle Eocene, Baja California SurENT#093; (Schweitzer et al., 2006b).

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 1893

Eurytium Stimpson, 1862

Eurytium sp. ENT#091;early Miocene, Chiapas; Pliocene, VeracruzENT#093; (Vega et al., 2009).

Panopeus H.Milne Edwards, 1834

Panopeus veintensisVega, Nyborg, Coutiño and Hernández-Monzón, 2008 ENT#091;early Eocene (Ypresian), ChiapasENT#093;.

XanthidaeMacLeay, 1838

Xanthidae incertae sedis

?HaydnellaMüller, 1984

?Haydnella sp. cf. H. steiningeriMüller, 1984 ENT#091;early Miocene, ChiapasENT#093; (Vega et al., 2009).

MegaxanthoVega, Feldmann, Garcia-Barrera, Filkorn, Pimentel and Avendaño, 2001b

Megaxantho zoqueVega, Feldmann, Garcia-Barrera, Filkorn, Pimentel and Avendaño, 2001b ENT#091;Maastrichtian, ChiapasENT#093; (Dietl and Vega, 2008).

Subsection THORACOTREMATAGuinot, 1977

Superfamily ?PINNOTHEROIDEADe Haan, 1833

?PinnotheridaeDe Haan, 1833

?PinnotherinaeDe Haan, 1833

ViapinnixaSchweitzer and Feldmann, 2001

Viapinnixa alvareziVega, Cosma, Coutiño, Feldmann, Nyborg, Schweitzer and Waugh, 2001a ENT#091;early Eocene (Ypresian), ChiapasENT#093; (Vega et al., 2008; Armstrong et al., 2009).

Viapinnixa perrilliataeVega, Nyborg, Fraaye and Espinosa, 2007a ENT#091;Paleocene (Selandian), CoahuilaENT#093; (Armstrong et al., 2009) (Fig. 13K).

Superfamily GRAPSOIDEAMacLeay, 1838

SesarmidaeDana, 1851c

Sesarmidae indet. ENT#091;Miocene (Aquitanian), ChiapasENT#093; (Serrano-Sánchez et al., 2016).

Brachyura incertae sedis

?Xanthosia Bell, 1863

? Xanthosia zoquiapensisFraaije, Vega, Van Bakel and Garibay-Romero, 2006 ENT#091;Campanian, GuerreroENT#093;.

RoemerusBishop, 1983b

Roemerus robustusBishop, 1983b ENT#091;Albian, ChiapasENT#093; (Vega et al., 2006a).

SOUTHERN AND CENTRAL FLORIDA, USA

Infraorder ANOMURAMacLeay, 1838

Superfamily GALATHEOIDEASamouelle, 1819

PorcellanidaeHaworth, 1825

Petrolisthes Stimpson, 1858

Petrolisthes myakkensisBishop and Portell, 1989 ENT#091;late Pliocene, Charlotte CountyENT#093; (Fig. 14A).

Figure 14 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 PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 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 bouvieriRathbun, 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 BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RanininaeDe Haan, 1839

LophoraninaFabiani, 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

EpialtidaeMacLeay, 1838

PisinaeDana, 1851b

Libinia Leach, 1815

Libinia sp. ENT#091;early-middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).

MithracidaeMacLeay, 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 suwanneeanaRathbun, 1935b ENT#091;late Eocene, Suwannee CountyENT#093;.

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus sp. ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).

CalappidaeDe 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 robertsiRoss, 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 brooksiRoss and Scolaro, 1964 ENT#091;late Eocene of Alachua, Marion, and Lafayette CountiesENT#093; (Portell, 2004) (Fig. 14D, E).

Calappilia calculosaRumsey, Klompmaker and Portell, 2016 ENT#091;late Eocene-early Oligocene, Suwannee CountyENT#093;.

Superfamily LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 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 PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 1838

Platylambrus Stimpson, 1871

Platylambrus charlottensis † (Rathbun, 1935b) ENT#091;early Pleistocene, Charlotte CountyENT#093; (Fig. 14G, H).

Superfamily CANCROIDEALatreille, 1802

CancridaeLatreille, 1802

CancrinaeLatreille, 1802

Cancer Latreille, 1802

Cancer irroratus Say, 1817 ENT#091;late Pleistocene-Holocene, Miami-Dade CountyENT#093; (Rathbun, 1935b).

Superfamily CHEIRAGONOIDEAOrtmann, 1893

CheiragonidaeOrtmann, 1893

MontezumellaRathbun, 1930

Montezumella microporosa † Portell and Collins, 2002 ENT#091;late Eocene, Alachua CountyENT#093; (Fig. 15F).

Figure 15 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 PORTUNOIDEARafinesque, 1815

OvalipidaeSpiridonov et al., 2014

Ovalipes Rathbun, 1898

Ovalipes stephensoniWilliams, 1976 ENT#091;early-middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).

PortunidaeRafinesque, 1815

PortuninaeRafinesque, 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 CARPILIOIDEAOrtmann, 1893

CarpiliidaeOrtmann, 1893

OcalinaRathbun, 1929

Ocalina floridanaRathbun, 1929 (type) ENT#091;late Eocene of Alachua, Levy, and Marion CountiesENT#093; (Rathbun, 1935b) (Fig. 15A-C).

ParaocalinaBeschin, 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 ERIPHIOIDEAMacLeay, 1838

MenippidaeOrtmann, 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 PILUMNOIDEASamouelle, 1819

PilumnidaeSamouelle, 1819

PilumninaeSamouelle, 1819

Pilumnus Leach, 1816

Pilumnus sp. ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).

Subsection THORACOTREMATAGuinot, 1977

Superfamily OCYPODOIDEARafinesque, 1815

OcypodidaeRafinesque, 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 PINNOTHEROIDEADe Haan, 1833

PinnotheridaeDe Haan, 1833

PinnothereliinaeAlcock, 1900a

Pinnixa White, 1846

Pinnixa sp. ENT#091;middle Pleistocene, Okeechobee CountyENT#093; (Agnew, 2001; Portell and Agnew, 2004).

Caribbean Islands & Bermuda

ANGUILLA

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa earleiWithers, 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 PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 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 PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Scylla De Haan, 1833

Scylla costataRathbun, 1919 (claws only) ENT#091;early Miocene, West side of Road BayENT#093; (as late Oligocene inWithers, 1924a).

PodophthalminaeDana, 1851d

PsygmophthalmusSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c

Psygmophthalmus bifurcatusCollins in Collins et al., 2009c ENT#091;early Miocene, Betty Hill QuarryENT#093;.

Superfamily XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 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 ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 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 BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

CalappidaeDe Haan, 1833

Mursia Leach inDesmarest, 1823

Mursia granulosaCollins and Donovan, 2002 ENT#091;late Oligocene, locality unknownENT#093; (Collins and Donovan, 2002, p. 145).

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Necronectes A. Milne-Edwards, 1881

Necronectes summusCollins and Donovan, 1995 ENT#091;late Oligocene, Nonsuch BayENT#093; (Fig. 16A).

Figure 16 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 THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 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;.

Gecarcinus Leach, 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 BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

CalappidaeDe Haan, 1833

Mursia Leach in Desmarest, 1823

Mursia creutzbergiCollins and Donovan, 2004 ENT#091;Miocene-Pliocene, Rooi TakiENT#093;.

BAHAMAS

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 1838

Gecarcinus Leach, 1814 ENT#091;in Leach, 1813-1815ENT#093;

Gecarcinus sp. ENT#091;late Holocene, San SalvadorENT#093; (Locatelli, 2013).

BARBADOS

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

EpialtidaeMacLeay, 1838

PisinaeDana, 1851b

Herbstia H.Milne Edwards, 1834

Herbstia exsertaCollins and Morris, 1976 ENT#091;middle to late Pleistocene, Coral RockENT#093;.

MithracidaeMacLeay, 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 PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

AtoportuninaeŠtevčić, 2005

Laleonectes Manning and Chace, 1990

Laleonectes vocans (A. Milne-Edwards, 1878) ENT#091;middle to late Pleistocene, Coral RockENT#093;.

PortuninaeRafinesque, 1815

Achelous De Haan, 1833

Achelous gibbesii (Stimpson, 1862) ENT#091;middle to late Pleistocene, Coral RockENT#093;.

Superfamily CARPILIOIDEAOrtmann, 1893

CarpiliidaeOrtmann, 1893

Carpilius Desmarest, 1823

Carpilius corallinus (Herbst, 1783) ENT#091;middle to late Pleistocene, Coral RockENT#093; (in Collins et al., 2009c).

Superfamily GONEPLACOIDEAMacLeay, 1838

ChasmocarcinidaeSerène, 1964

ChasmocarcininaeSerène, 1964

FalconoplaxVan Straelen, 1933b

Falconoplax bicarinellaCollins and Morris, 1976 ENT#091;early-middle Eocene Scotland Beds, SpaENT#093;.

Superfamily HEXAPODOIDEAMiers, 1886

HexapodidaeMiers, 1886

PalaeopinnixaVía Boada, 1966

Palaeopinnixa perornataCollins and Morris, 1976 (type) ENT#091;early-middle Eocene Scotland Beds, SpaENT#093;.

Superfamily PILUMNOIDEASamouelle, 1819

PilumnidaeSamouelle, 1819

PilumninaeSamouelle, 1819

Pilumnus Leach, 1816

Pilumnus sp. (claw fragment) ENT#091;Pleistocene, ClaphamENT#093; (Collins and Morris, 1976).

Superfamily XANTHOIDEAMacLeay, 1838

XanthidaeMacLeay, 1838

ActaeinaeAlcock, 1898

Paractaea Guinot, 1969

Paractaea nodosa (Stimpson, 1860) ENT#091;Pleistocene, Coral RockENT#093; (as Actaea rufopunctata inCollins and Morris, 1976).

BERMUDA

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 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 BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Superfamily CALAPPOIDEADe Haan, 1833

CalappidaeDe Haan, 1833

Calappilia † A.Milne-Edwards, 1873

Calappilia bonairensisVan Straelen, 1933a ENT#091;middle to late Eocene (Lutetian-Priabonian), southwest of Seroe MontagneENT#093;.

Superfamily CHEIRAGONOIDEAOrtmann, 1893

CheiragonidaeOrtmann, 1893

MontezumellaRathbun, 1930

Montezumella rutteniVan Straelen, 1933a ENT#091;middle to late Eocene (Lutetian-Priabonian), southwest of Seroe MontagneENT#093; (also reported inCollins and Donovan, 2005).

Superfamily CARPILIOIDEAOrtmann, 1893

CarpiliidaeOrtmann, 1893

OcalinaRathbun, 1929

Ocalina sublevisCollins and Donovan, 2005 ENT#091;middle to late Eocene (Lutetian-Priabonian), southwest of Seroe MontagneENT#093;.

CUBA

Infraorder ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

PaguridaeLatreille, 1802

Paguridae indet. ENT#091;late Pleistocene, US Guantanamo Bay Naval StationENT#093; (Collins et al., 2009c).

Infraorder BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RanininaeDe Haan, 1839

VegaraninaVan 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).

RaninoidinaeLőrenthey 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

EpialtidaeMacLeay, 1838

PisinaeDana, 1851b

Libinia Leach, 1815

Libinia sp. ENT#091;?Oligocene-Miocene, locality unknownENT#093; (Varela, 2013).

MithracidaeMacLeay, 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 CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus sp. ENT#091;early Miocene, Matanzas ProvinceENT#093; (in Varela and Rojas-Consuegra, 2011b).

EriosachilaBlow and Manning, 1996

Eriosachila cubaensisVarela and Rojas-Consuegra, 2011a ENT#091;early Miocene, Matanzas ProvinceENT#093;.

CalappidaeDe Haan, 1833

Calappidae indet. ENT#091;late Pleistocene, US Guantanamo Bay Naval StationENT#093; (Collins et al., 2009c).

Superfamily LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 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 PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 1838

Spinolambrus Tan andNg, 2007

Spinolambrus lazaroiVarela, 2013 ENT#091;early Miocene, Matanzas ProvinceENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Necronectes † A.Milne-Edwards, 1881

Necronectes collinsiSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c ENT#091;early Miocene, Matanzas ProvinceENT#093; (Varela and Rojas-Consuegra, 2011b).

PodophthalminaeDana, 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 andSchweitzer, 2011

Paraeuphylax cubaensis † (type) Varela and Schweitzer, 2011 ENT#091;early Miocene, Matanzas ProvinceENT#093;.

PortuninaeRafinesque, 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 oblongusRathbun, 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 ERIPHIOIDEAMacLeay, 1838

EriphiidaeMacLeay, 1838

Eriphia Latreille, 1817

Eriphia sp. ENT#091;early Miocene, Sancti Spíritus ProvinceENT#093; (Varela, 2013).

Superfamily XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

Panopeidae indet. ENT#091;late Pleistocene, US Guantanamo Bay Naval StationENT#093; (Collins et al., 2009c).

PanopeinaeOrtmann, 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 THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 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 BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

NotopodinaeSerène 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).

RaninoidinaeLőrenthey 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

MithracidaeMacLeay, 1838

Maguimithrax Klompmaker et al., 2015

Maguimithrax spinosissimus (Lamarck, 1818) ENT#091;Pliocene, Salina Sint MichaelENT#093; (Stepp, 2014; Klompmaker et al., 2015).

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus lineatinusCollins and Todd in Todd and Collins, 2005 ENT#091;Pliocene, Salina Saint MichielENT#093; (Stepp, 2014).

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa galloidesStimpson, 1859 ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014).

Superfamily LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 1871

Myropsis Stimpson, 1871

Myropsis quinquespinosaStimpson, 1871 ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014)

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

PortuninaeRafinesque, 1815

Achelous De Haan, 1833

Achelous sebae (H. Milne Edwards, 1834) ENT#091;Pliocene, Salina Sint MichielENT#093; (Stepp, 2014).

DOMINICAN REPUBLIC

Infraorder ANOMURAMacLeay, 1838

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 1892

Dardanus Paul’son, 1875

Dardanus squamatusCollins in Collins et al., 2009c ENT#091;late Miocene-early Pliocene, Río Gurabo and Río MaoENT#093;.

Petrochirus Stimpson, 1858

Petrochirus inequalisRathbun, 1919 ENT#091;Pleistocene, Rio Gurabo and Santo DomingoENT#093; (Collins et al., 2009c).

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus guraboensisCollins in Collins et al., 2009c ENT#091;late Miocene, Rio Gurabo, Rio Cana and Rio MaoENT#093;.

CalappidaeDe 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 LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 1871

Persephona Leach, 1817

Persephona prepunctataRathbun, 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 PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 1838

Mesorhoea Stimpson, 1871

Mesorhoea mauryaeRathbun, 1919 ENT#091;late Miocene-early Pliocene, Río Cana and Santo DomingoENT#093; (Collins et al., 2009c).

Platylambrus Stimpson, 1871

Platylambrus obscuraRathbun, 1919 ENT#091;Miocene, Santo DomingoENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Scylla De Haan, 1833

Scylla costataRathbun, 1919 ENT#091;Miocene, valley of Yaque del Norte RiverENT#093; ENT#091;Note 1ENT#093;.

PodophthalminaeDana, 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;.

PsygmophthalmusSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c

Psygmophthalmus bifurcatusCollins in Collins et al., 2009c ENT#091;age uncertain, Rio CanaENT#093;.

SandomingiaRathbun, 1919

Sandomingia yaquiensisRathbun, 1919 ENT#091;late Miocene-early Pliocene, Río Cana and Rio Yaque del NorteENT#093; (Collins et al., 2009c) ENT#091;Note 1ENT#093;.

PortuninaeRafinesque, 1815

Callinectes Stimpson, 1862

Callinectes declivisRathbun, 1918 (fingers only) ENT#091;late Miocene, Rio Gurabo and Rio Mao Santo DomingoENT#093; (inCollins et al., 2009c).

Portunus Weber, 1795

Portunus gabbiRathbun, 1919 ENT#091;Miocene, valley of Yaque del Norte RiverENT#093; ENT#091;Note 1ENT#093;.

Portunus oblongusRathbun, 1920b ENT#091;late Miocene-early Pliocene, Río CanaENT#093;.

Portunus tenuisRathbun, 1919 (type) (claws only) ENT#091;Miocene of Yaque Valley, Santo DomingoENT#093;.

RathbunitesSchweitzer, 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 CARPILIOIDEAOrtmann, 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 PILUMNOIDEASamouelle, 1819

PilumnidaeSamouelle, 1819

PilumninaeSamouelle, 1819

Pilumnus Leach, 1816

Pilumnus subequusRathbun, 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 XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

CalappidaeDe 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 ERIPHIOIDEAMacLeay, 1838

PlatyxanthidaeGuinot, 1977

Platyxanthus A.Milne-Edwards, 1863

Platyxanthus sp. ENT#091;middle Miocene, CarriacouENT#093; (Collins et al., 2009c).

HAITI

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Eubrachyuran indet. (claw) (as ?Zanthopsis indet. inRathbun, 1923a) ENT#091;Eocene, Artibonite departmentENT#093; ENT#091;Note 2ENT#093;.

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

MithracidaeMacLeay, 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 PARTHENOPOIDEAMacLeay, 1838

?Parthenopoid indet. (claw fragment) ENT#091;early Miocene, Centre departmentENT#093; ENT#091;Note 2ENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

PortuninaeRafinesque, 1815

Portunus Weber, 1795

Portunus haitensisRathbun, 1923a ENT#091;early Miocene, Centre departmentENT#093; ENT#091;Note 1ENT#093;.

Superfamily XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 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 ANOMURAMacLeay, 1838

Superfamily GALATHEOIDEASamouelle, 1819

PorcellanidaeHaworth, 1825

Petrolisthes Stimpson, 1858

Petrolisthes sp. ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).

Superfamily HIPPOIDEALatreille, 1825

AlbuneidaeStimpson. 1858

Albunea Weber, 1975

Albunea sp. ENT#091;late Pleistocene, parish of St. Ann) (Morris, 1993).

Superfamily PAGUROIDEALatreille, 1802

DiogenidaeOrtmann, 1892

Dardanus Paul’son, 1875

Dardanus portmorantensisCollins 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 BRACHYURALatreille, 1802

Superfamily HOMOLODROMIOIDEAAlcock, 1900b

Goniodromitidae †Beurlen, 1932

TrechmanniusCollins and Donovan, 2006

Trechmannius circularisCollins and Donovan, 2006 (type) ENT#091;early Paleocene, parish of PortlandENT#093;.

Superfamily DROMIOIDEADe Haan, 1833

DynomenidaeOrtmann, 1892

Dynomene Desmarest, 1823

Dynomene variabilis † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.

KromtitisMüller, 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

CretacoraninaMertin, 1941

Cretacoranina trechmanni † (Withers, 1927, as Ranina trechmanni) ENT#091;Maastrichtian, parish of St. JamesENT#093; (Morris, 1993).

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RaninoidinaeLőrenthey 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily MAJOIDEASamouelle, 1819

EpialtidaeMacLeay, 1838

PisinaeDana, 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).

MithracidaeMacLeay, 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 arawakumKlompmaker 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 windsoraeKlompmaker 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;.

OregoniidaeGarth, 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 CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus praecoxCollins, 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).

CalappidaeDe 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 LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 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

DuncanitrixSchweitzer, Dworschak and Martin, 2011ENT#091;=Duncania Portell andCollins, 2004ENT#093;

Duncanitrix jamaicensis † (Portell and Collins, 2004) ENT#091;early Miocene, parish of TrelawnyENT#093;.

Superfamily PARTHENOPOIDEAMacLeay, 1838

ParthenopidaeMacLeay, 1838

ParthenopinaeMacLeay, 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 CANCROIDEALatreille, 1802

CancridaeLatreille, 1802

CancrinaeLatreille, 1802

Cancer Linnaeus, 1758

Cancer sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).

Superfamily PORTUNOIDEARafinesque, 1815

Carcineretidae †Beurlen, 1930

CarcineretesWithers, 1922

Carcineretes woolacottiWithers, 1922 ENT#091;Maastrichtian, parishes of Clarendon and St. JamesENT#093;.

OvalipidaeSpiridonov et al., 2014

Ovalipes Rathbun, 1898

Ovalipes sp. ENT#091;late Pliocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).

PortunidaeRafinesque, 1815

AtoportuninaeŠtevčić, 2005

Laleonectes Manning and Chace, 1990

Laleonectes vocans (A. Milne-Edwards, 1878) ENT#091;late Pleistocene, parish of St. ThomasENT#093;.

PodophthalminaeDana, 1851d

Euphylax Stimpson, 1862

Euphylax fortispinosusCollins, Donovan, Lindsay and Simpson, 2001 ENT#091;early Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009c).

PortuninaeRafinesque, 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 jamaicensisWithers, 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 andCollins, 2004

Pseudoachelous schindleri † Portell and Collins, 2004 (type) ENT#091;early Miocene, parish of TrelawnyENT#093;.

Superfamily CARPILIOIDEAOrtmann, 1893

CarpiliidaeOrtmann, 1893

Carpilius Desmarest, 1823

Carpilius corallinus (Herbst, 1783) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins et al., 2009a).

OcalinaRathbun, 1929

Ocalina haldixoniCollins 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;.

PalaeoxanthopsisBeurlen, 1958b

Palaeoxanthopsis sp. † (as ?Paranecrocarcinus sp. inMorris, 1993, fig. 1.6) ENT#091;?Maastrichian, unknownENT#093; ENT#091;Note 2ENT#093;.

Superfamily DAIROIDEASerène, 1965

DairidaeSerène, 1965

Daira De Haan, 1833

Daira vulgaris † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093; (Fig. 16E)

Superfamily ERIPHIOIDEAMacLeay, 1838

EriphiidaeMacLeay, 1838

Eriphia Latreille, 1817

Eriphia gonagra xaymacaensisCollins 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 PILUMNOIDEASamouelle, 1819

PilumnidaeSamouelle, 1819

PilumninaeSamouelle, 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 TRAPEZIOIDEAMiers, 1886

TrapeziidaeMiers, 1886

Trapezia Latreille, 1828

Trapezia prisca † Portell and Collins, 2004 ENT#091;early Miocene, parish of TrelawnyENT#093;.

Superfamily XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 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.).

PseudorhombilidaeAlcock, 1900a

Nanoplax Guinot, 1967

Nanoplax xanthiformis (A. Milne-Edwards, 1880b) ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins and Portell, 1998).

XanthidaeMacLeay, 1838

ActaeinaeAlcock, 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 andCollins, 2004

Actaeops frontalis † Portell and Collins, 2004 (type) ENT#091;early Miocene, parish of TrelawnyENT#093;.

ChlorodiellinaeNg 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;.

XanthinaeMacLeay, 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 rathbunaeWithers, 1924b ENT#091;middle Eocene (Lutetian), parish of HanoverENT#093; (Morris, 1993).

Subsection THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 1838

Cardisoma Latreille, 1828

Cardisoma guanhumi Latreille, 1828 ENT#091;late Pleistocene, parish of St. ThomasENT#093; (Collins and Donovan, 1998; 2012).

GrapsidaeMacLeay, 1838

Pachygrapsus Randall, 1840

Pachygrapsus sp. ENT#091;late Pleistocene, parish of St. AnnENT#093; (Morris, 1993).

SesarmidaeDana, 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 OCYPODOIDEARafinesque, 1815

OcypodidaeRafinesque, 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 BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RanininaeDe Haan, 1839

VegaraninaVan 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 EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa pavimentaSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c ENT#091;early Miocene, San SebastiánENT#093;.

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 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 costataRathbun, 1919 (Miocene, San Sebastián and YaucoENT#093; (Schweitzer et al., 2008).

PodophthalminaeDana, 1851d

PsygmophthalmusSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c

Psygmophthalmus laresSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c (type) ENT#091;early Miocene, San SebastiánENT#093;.

PortuninaeRafinesque, 1815

Portunus Weber, 1795

Portunus oblongusRathbun, 1920b ENT#091;Miocene, uncertainENT#093; (Gordon, 1966; Schweitzer et al., 2006c).

Portunus yaucoensisSchweitzer, Iturralde-Vinent, Hetler and Velez-Juarbe, 2006c ENT#091;early Oligocene (Rupelian), YaucoENT#093;.

Superfamily CARPILIOIDEAOrtmann, 1893

Palaeoxanthopsidae †Schweitzer, 2003

PalaeoxanthopsisBeurlen, 1958b

Palaeoxanthopsis tylotus † Schweitzer, Velez-Juarbe, Martinez, Collmar Hull, Feldmann and Santos, 2008 ENT#091;late Maastrichtian, Sabana GrandeENT#093;.

Superfamily XANTHOIDEAMacLeay, 1838

PanopeidaeOrtmann, 1893

PanopeinaeOrtmann, 1893

Eurytium Stimpson, 1859

Eurytium granulosumSchweitzer, Velez-Juarbe, Martinez, Hull, Feldmann and Santos, 2008 ENT#091;Miocene, near PonceENT#093; (Schweitzer et al., 2008).

Subsection THORACOTREMATAGuinot, 1977

Superfamily GRAPSOIDEAMacLeay, 1838

GecarcinidaeMacLeay, 1838

Cardisoma Latreille, 1828

Cardisoma guanhumiLatreille, 1828 ENT#091;Pleistocene, UtuadoENT#093; (Schweitzer et al., 2008).

SAINT BARTHÉLEMY (ST. BARTS)

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

EriosachilaBlow and Manning, 1996

Eriosachila bartholomaensis † (Rathbun, 1919, as Zanthopsis bartholomaensis) ENT#091;Eocene, St. BartholomewENT#093;.

SAINT MARTIN

Infraorder BRACHYURALatreille, 1802

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

CalappidaeDe Haan, 1833

TutusCollins in Collins et al., 2009c

Tutus granulosusCollins in Collins et al., 2009c ENT#091;Miocene of Leeward Islands, Tintamare IslandENT#093;.

TRINIDAD

Infraorder BRACHYURALatreille, 1802

Superfamily RANINOIDEADe Haan, 1839

RaninidaeDe Haan, 1839

RanininaeDe Haan, 1839

LophoraninaFabiani, 1910

Lophoranina porifera † (Woodward, 1866) ENT#091;‘Tertiary’ indet, San FernandoENT#093;.

Section EUBRACHYURASaint Laurent, 1980

Subsection HETEROTREMATAGuinot, 1977

Superfamily CALAPPOIDEADe Haan, 1833

AethridaeDana, 1851d

Hepatus Latreille, 1802

Hepatus nodosusCollins and Morris, 1976 ENT#091;middle Miocene, Manzanilla BayENT#093;.

CalappidaeDe Haan, 1833

Calappa Weber, 1795

Calappa cuspidata † (Guppy, 1909, as Ranina cuspidata) ENT#091;Oligocene, Tamana DistrictENT#093; (Pasini and Garassino, 2017).

Superfamily LEUCOSIOIDEASamouelle, 1819

LeucosiidaeSamouelle, 1819

EbaliinaeStimpson, 1871

Persephona Leach, 1817

Persephona punctata (Linnaeus, 1758) ENT#091;Miocene-Pliocene, Manzanilla BayENT#093; (Collins and Morris, 1976).

Superfamily PORTUNOIDEARafinesque, 1815

PortunidaeRafinesque, 1815

NecronectinaeGlaessner, 1928

Necronectes † A.Milne-Edwards, 1881

Necronectes proavitus † (Rathbun, 1918) ENT#091;middle Miocene, near Caparo Saca MantecaENT#093; (Collins and Morris, 1976).

PortuninaeRafinesque, 1815

Portunus Weber, 1795

Portunus oblongusRathbun, 1920b ENT#091;middle Miocene, Manzanilla Bay and MontserratENT#093; (Collins and Morris, 1976).

Superfamily HEXAPODOIDEAMiers, 1886

HexapodidaeMiers, 1886

PalaeopinnixaVía Boada, 1966

Palaeopinnixa intermedia † (Collins and Morris, 1976, as Thaumastoplax intermedia) ENT#091;middle Miocene, MontserratENT#093;

Palaeopinnixa perornataCollins 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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1This article is part of the tribute offered by the Brazilian Crustacean Society in memoriam of Michael Türkay for his outstanding contribution to Carcinology.

2Guest Editor: Célio Magalhães

Received: February 16, 2017; Accepted: June 26, 2017

Corresponding author: Javier Luque. luque@ualberta.ca

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