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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.