Journal of Mammalian Evolution

, Volume 24, Issue 1, pp 71–90 | Cite as

Fossil Cetaceans (Mammalia, Cetacea) from the Neogene of Colombia and Venezuela

  • Gabriel Aguirre-FernándezEmail author
  • Jorge D. Carrillo-Briceño
  • Rodolfo Sánchez
  • Eli Amson
  • Marcelo R. Sánchez-Villagra
Original Paper


There are significant geographic gaps in our knowledge of marine mammal evolution because most fossils have been found and described from Northern Hemisphere localities and a few other high-latitude areas in the Southern Hemisphere. Here, we describe fossil cetacean remains from five geological units in the South American tropics (Urumaco, Codore, Castilletes, Cantaure, and Querales formations) generally representing marginal marine depositional environments (estuaries, deltas, and tidal flats). While fossil cetaceans from Venezuelan Neogene localities have been previously studied, this paper includes the first descriptions of fossil cetaceans from Colombia, including a diverse assemblage of mysticetes and odontocetes. We identified and provisionally referred fragmentary remains to the iniid Ischyrorhynchus vanbenedeni and to the platanistid Zarhachis flagellator. The latter suggests the presence of Platanistidae in the eastern coast of South America during the early-middle Miocene, representing the second record of Platanistidae in South America and the first record of Platanistidae in eastern South America. Other less-diagnostic specimens are characterized by features commonly seen in longirostrine odontocetes such as Iniidae, Platanistidae, Pontoporiidae, Lipotidae, Eoplatanistidae, and Squalodelphinidae.


Odontoceti Mysticeti Miocene Cocinetas and Falcón Basins South American Tropics 



This work greatly benefited from the suggestions of the reviewers (Giovanni Bianucci and Olivier Lambert) and the editors (François Pujos, Pierre-Olivier Antoine, and John Wible). We thank Carlos Jaramillo (STRI), Analía Forasiepi (CCT-CONICET-Mendoza), and Yoshihiro Tanaka (Numata Fossil Museum) for reading through and commenting on earlier versions of this manuscript. Juan David Carrillo (University of Zurich), Mary Parra and Fredy Parra (CIP) are thanked for fossil preparation. Loïc Costeur (NMB), Mary Parra (STRI), and Jaime Escobar (Universidad del Norte) kindly provided access to specimens under their care. Thanks to Andrés Reyes Cespedes (Universidad Experimental Francisco de Miranda) for help in collections. Thanks to Michael Knappertbusch (NMB) for guidance with micropaleontological analysis and access to laboratory and equipment. Austin Hendy (LACM) and Gustavo Ballen (STRI) kindly provided contextual information regarding the fieldwork in which the MUN-STRI specimens were collected. Juan David Carrillo, Ashley Latimer, and Torsten Scheyer (University of Zurich) are thanked for useful discussions; Juan David Carrillo provided measurements of AMU-CURS-219. GAF wishes to thank Jairo Carrillo (Colombia)and his family for their hospitality and logistical aid. Thanks to Scott Hartman for producing the silhouette used in Fig. 1 and PhyloPic for making it available.

We thank the Instituto del Patrimonio Cultural de Venezuela and the Alcaldía Bolivariana de Urumaco, for the authorization and permissions to collect and study the fossil material from Venezuela. The Smithsonian Institution, the National Geographic Society, Universidad del Norte, the Anders Foundation, Gregory D. and Jennifer Walston Johnson, and the National Science Foundation (grant EAR 0957679) helped to support fieldwork. Thanks to Carlos Rosero for managing all the logistics in the field in Colombia, Liliana Londoño and Maria Ines Barreto for administrative and logistic support. Thanks to the Wayuú Community for allowing access to their lands and for support during fieldwork, the Colombian National Police (Castilletes base), and all the members of the field teams over the past four years.

This project was funded by Swiss National Science Foundation grant 31003A_149605 to MRS-V. GAF was supported by Fonds für Lehre und Forschung from the Naturhistorisches Museum Basel. EA was supported by a Smithsonian Tropical Research Institute postdoctoral fellowship grant.

Supplementary material

10914_2016_9353_MOESM1_ESM.pdf (1.6 mb)
Supplementary data The Online Resource 1 includes unlabeled specimen photographs. Additional field data linked to MUN-STRI specimens can be downloaded from the Smithsonian Tropical Research Institute Geological Sample Database webpage ( using either the locality (e.g., “490006”) or the sample (e.g., “37698”) numbers. (PDF 1656 kb)


  1. Aguilera OA (2004) Tesoros paleontológicos de Venezuela: Urumaco, patrimonio natural de la humanidad. Editorial Arte, VenezuelaGoogle Scholar
  2. Aguilera OA, García L, Cozzuol MA (2008) Giant-toothed white sharks and cetacean trophic interaction from the Pliocene Caribbean Paraguaná Formation. Paläontol Z 82:204–208. doi:  10.1007/BF02988410 CrossRefGoogle Scholar
  3. Aguilera OA, Moraes-Santos H, Costa S, Ohe F, Jaramillo C, Nogueira A (2013) Ariid sea catfishes from the coeval Pirabas (northeastern Brazil), Cantaure, Castillo (Northwestern Venezuela), and Castilletes (north Colombia) formations (early Miocene), with description of three new species. Swiss J Palaeontol 132:45–68CrossRefGoogle Scholar
  4. Ameghino F (1891) Caracteres diagnósticos de cincuenta especies nuevas de mamíferos fósiles argentinos. Rev Argent Hist Nat 1:129–167Google Scholar
  5. Barnes LG (1985) Fossil pontoporiid dolphins (Mammalia: Cetacea) from the Pacific coast of North America. Contrib Sci 363:1–34Google Scholar
  6. Barnes LG (2006) A phylogenetic analysis of the superfamily Platanistoidea (Mammalia, Cetacea, Odontoceti). Beitr Palaeontol 30:25–42Google Scholar
  7. Bengtson P (1988) Open nomenclature. Palaeontology 31:223–227Google Scholar
  8. Best RC, da Silva VMF (1993) Inia geoffrensis. Mammal Species 1–8Google Scholar
  9. Bianucci G (2005) Arimidelphis sorbinii a new killer whale-like dolphin from the Pliocene of the Marecchia River (central eastern Italy) and a phyogenetic analysis of the Orcininae (Cetacea: Odonticeti). Riv Ital Paleontol S 111:329–344Google Scholar
  10. Bianucci G, Di Celma C, Landini W, Post K, Tinelli C, Muizon C de, Gariboldi K, Malinverno E, Cantalamessa G, Gioncada A, Collareta A, Gismondi R-S, Varas-Malca R, Urbina M, Lambert O (2016) Distribution of fossil marine vertebrates in Cerro Colorado, the type locality of the giant raptorial sperm whale Livyatan melvillei (Miocene, Pisco Formation, Peru). J Maps 12:543–557. doi:  10.1080/17445647.2015.1048315
  11. Bianucci G, Lambert O, Post K (2010) High concentration of long-snouted beaked whales (genus Messapicetus) from the Miocene of Peru. Palaeontology 53:1077–1098. doi:  10.1111/j.1475-4983.2010.00995.x CrossRefGoogle Scholar
  12. Bianucci G, Lambert O, Salas-Gismondi R, Tejada J, Pujos F, Urbina M, Antoine P-O (2013) A Miocene relative of the Ganges River dolphin (Odontoceti, Platanistidae) from the Amazonian Basin. J Vertebr Paleontol 33:741–745. doi:  10.1080/02724634.2013.734888 CrossRefGoogle Scholar
  13. Bianucci G, Landini W (2002) Change in diversity, ecological significance and biogeographical relationships of the Mediterranean Miocene toothed whale fauna. Geobios 24:19–28Google Scholar
  14. Bianucci G, Landini W, Valleri G, Ragaini L, Varola A (2005) First cetacean fossil records from Ecuador, collected from the Miocene of Esmeraldas Province. Riv Ital Paleontol S 111:345–350Google Scholar
  15. Bianucci G, Urbina M, Lambert O (2015) A new record of Notocetus vanbenedeni (Squalodelphinidae, Odontoceti, Cetacea) from the early Miocene of Peru. C R Palevol 14:5–13. doi:  10.1016/j.crpv.2014.08.003 CrossRefGoogle Scholar
  16. Buchholtz EA (2001) Vertebral osteology and swimming style in living and fossil whales (Order: Cetacea). J Zool 253:175–190CrossRefGoogle Scholar
  17. Buchholtz EA, Schur SA (2004) Vertebral osteology in Delphinidae (Cetacea). Zool J Linn Soc 140:383–401CrossRefGoogle Scholar
  18. Buono MR, Cozzuol MA (2013) A new beaked whale (Cetacea, Odontoceti) from the late Miocene of Patagonia, Argentina. J Vertebr Paleontol 33:986–997. doi:  10.1080/02724634.2013.752377 CrossRefGoogle Scholar
  19. Buono MR, Viglino M, Cozzuol MA, Lucero S (2016) Descifrando la historia evolutiva de los Neoceti (Mammalia: Cetacea): aportes del registro fósil del Neógeno del Atlántico Sudoccidental. In: Agnolin FL, Lio GL, Brissón Egli F, Chimento NR, Novas FE (ed) Historia Evolutiva y Paleobiogeográfica de los Vertebrados de América del Sur, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,” Buenos Aires, pp 323–334Google Scholar
  20. Cabrera A (1926) Cetáceos fósiles del Museo de la Plata. Rev Mus La Plata 24:363–411Google Scholar
  21. Carrillo JD, Forasiepi A, Jaramillo C, Sánchez-Villagra MR (2015) Neotropical mammal diversity and the Great American Biotic Interchange: spatial and temporal variation in South America’s fossil record. Front Genet 5. doi:  10.3389/fgene.2014.00451
  22. Carrillo-Briceño JD, Aguilera OA, De Gracia C, Aguirre-Fernández G, Kindlimann R, Sánchez-Villagra MR (2016) An Early Neogene Elasmobranch fauna from the southern Caribbean (western Venezuela). Palaeontol Electron 19.2.28AGoogle Scholar
  23. Carrillo-Briceño JD, Maxwell E, Aguilera OA, Sánchez R, Sánchez-Villagra MR (2015) Sawfishes and other elasmobranch assemblages from the Mio-Pliocene of the south Caribbean (Urumaco sequence, northwestern Venezuela). PLoS ONE 10:e0139230. doi:  10.1371/journal.pone.0139230 CrossRefPubMedPubMedCentralGoogle Scholar
  24. Cope ED (1868) Second contribution to the history of the Vertebrata of the Miocene period of the United States. Proc Acad Nat Sci Philadelphia 20:184–194. doi:  10.2307/4059884 Google Scholar
  25. Cope ED (1869) Third contribution to the fauna of the Miocene period of the United States. Proc Acad Nat Sci Philadelphia 21:6–12. doi:  10.2307/4060091 Google Scholar
  26. Cozzuol MA (1988) Una nueva especie de Saurodelphis Burmeister, 1891 (Cetacea: Iniidae) del “Mesopotamiense” (Mioceno tardío-Plioceno temprano) de la provincia de Entre Ríos, Argentina. Ameghiniana 25:39–45Google Scholar
  27. Cozzuol MA (1996) The record of the aquatic mammals in southern South America. Münchner Geowiss Abh 30:321–342Google Scholar
  28. Cozzuol MA (2006) The Acre vertebrate fauna: age, diversity, and geography. J South Am Earth Sci 21:185–203. doi:  10.1016/j.jsames.2006.03.005 CrossRefGoogle Scholar
  29. Cozzuol MA (2010) Fossil record and the evolutionary history of Inioidea. In: Ruiz-García M, Shostell JM (eds) Biology, Evolution and Conservation of River Dolphins within South America and Asia. Nova Science Publishers, New York, pp 193–217Google Scholar
  30. Cozzuol MA, Aguilera OA (2008) Cetacean remains from the Neogene of northwestern Venezuela. Paläontol Z 82:196–203.  10.1007/BF02988409 CrossRefGoogle Scholar
  31. Díaz de Gamero ML (1974) Microfauna y edad de la Formación Cantaure, Península de Paraguaná, Venezuela. Bol Inf Asoc Venez Geol Min Petr 17:41–47Google Scholar
  32. Díaz de Gamero ML (1989) El Mioceno temprano y medio de Falcón septentrional. GEOS 29:25–35Google Scholar
  33. Doyle P, Mather AE, Bennett MR, Bussell MA (1996) Miocene barnacle assemblages from southern Spain and their palaeoenvironmental significance. Lethaia 29:267–274. doi:  10.1111/j.1502-3931.1996.tb01659.x CrossRefGoogle Scholar
  34. Fordyce RE (1994) Waipatia maerewhenua, new genus and new species (Waipatiidae, new family), an archaic late Oligocene dolphin (Cetacea: Odontoceti: Platanistoidea) from New Zealand. In: Berta A, Deméré T (eds) Contributions in Marine Mammal Paleontology Honoring Frank C. Whitmore, Jr. Proc San Diego Soc of Nat Hist 29, pp 147–176Google Scholar
  35. Fordyce RE (2009) Fossil sites, noted. In: Perrin WF, Würsig B, Thewissen JGM (eds) Encyclopedia of Marine Mammals, 2nd ed. Academic Press, San Diego, pp 459–465CrossRefGoogle Scholar
  36. Fordyce RE, Muizon C de (2001) Evolutionary history of cetaceans: a review. In: Mazin J-M, Buffrenil V de (eds) Secondary Adaptation of Tetrapods to Life in Water. Proceedings of the International Meeting in Poitiers, 1996, Pfeil, Munich, pp 169–233Google Scholar
  37. Geisler JH, Godfrey SJ, Lambert O (2012) A new genus and species of late Miocene inioid (Cetacea, Odontoceti) from the Meherrin River, North Carolina, U.S.A. J Vertebr Paleontol 32:198–211. doi:  10.1080/02724634.2012.629016 CrossRefGoogle Scholar
  38. Geisler JH, McGowen MR, Yang G, Gatesy J (2011) A supermatrix analysis of genomic, morphological, and paleontological data from crown Cetacea. BMC Evol Biol 11:1–33Google Scholar
  39. Gibson-Smith W (1979) The genus Arcinella (Mollusca: Bivalvia) in Venezuela and some associated faunas. GEOS 24:11–32Google Scholar
  40. Gibson-Smith J, Gibson-Smith W (1983) Neogene melogenid gastropods from the Paraguaná Peninsula, Venezuela. Eclogae geol Helv 76:719–728Google Scholar
  41. Griffiths N, Müller W, Johnson KG, Aguilera OA (2013) Evaluation of the effect of diagenetic cements on element/Ca ratios in aragonitic early Miocene (~16 Ma) Caribbean corals: Implications for ‘deep-time’ palaeo-environmental reconstructions. Palaeogeogr Palaeoclimatol Palaeoecol 369:185–200. doi:  10.1016/j.palaeo.2012.10.018 CrossRefGoogle Scholar
  42. Gutstein CS, Cozzuol MA, Vargas AO, Suárez ME, Schultz CL, Rubilar-Rogers D (2009) Patterns of skull variation of Brachydelphis (Cetacea, Odontoceti) from the Neogene of the southeastern Pacific. J Mammal 90:504–519CrossRefGoogle Scholar
  43. Hambalek N, Rull V, Digiacomo E de, Díaz de Gamero M (1994) Evolución paleoecológica y paleoambiental de la secuencia del Neógeno en el surco de Urumaco. Estudio palinológico y litológico. Bol Soc Venezolana Geol 191:7–19Google Scholar
  44. Hendy AJW, Jones DS, Moreno F, Zapata V, Jaramillo C (2015) Neogene molluscs, shallow marine paleoenvironments, and chronostratigraphy of the Guajira Peninsula, Colombia. Swiss J Palaeontol 134:45–75. doi:  10.1007/s13358-015-0074-1 CrossRefGoogle Scholar
  45. Hodson F (1926) Venezuelan and Caribbean turritellas, with a list of type stratigraphic localities. Bull Am Paleontol 11:173–220Google Scholar
  46. Hunter VF, Bartok P (1974) The age and correlation of the Tertiary sediments of the Paraguaná Península, Venezuela. Bol Inf Asoc Venezolana Geol Min Petr 17:143–154Google Scholar
  47. Jaramillo C, Moreno F, Hendy AJW, Sánchez-Villagra MR, Marty D (2015) Preface: La Guajira, Colombia: a new window into the Cenozoic neotropical biodiversity and the Great American Biotic Interchange. Swiss J Palaeontol 134:1–4. doi:  10.1007/s13358-015-0075-0 CrossRefGoogle Scholar
  48. Jung P (1965) Miocene Mollusca from the Paraguaná Peninsula, Venezuela. Bull Am Paleontol 49:389–607Google Scholar
  49. Kellogg AR (1924) A fossil porpoise from the Calvert Formation of Maryland. Proc US Natl Mus 63:1–39CrossRefGoogle Scholar
  50. Kellogg AR (1927) Kentriodon pernix, a Miocene porpoise from Maryland. Proc US Natl Mus 69:1–55CrossRefGoogle Scholar
  51. Lambert O (2005) Phylogenetic affinities of the long-snouted dolphin Eurhinodelphis (Cetacea, Odontoceti) from the Miocene of Antwerp, Belgium. Palaeontology 48:653–679CrossRefGoogle Scholar
  52. Lambert O, Estevens M, Smith R (2005) A new kentriodontine dolphin from the middle Miocene of Portugal. Acta Palaeontol Pol 50:239–248Google Scholar
  53. Lambert O, Muizon C de, Bianucci G (2015) A new archaic homodont toothed cetacean (Mammalia, Cetacea, Odontoceti) from the early Miocene of Peru. Geodiversitas 37:79–108. doi:  10.5252/g2015n1a4
  54. Leduc RG, Perrin WF, Dizon AE (1999) Phylogenetic relationships among the delphinid cetaceans based on full cytochrome b sequences. Mar Mammal Sci 15:619–648CrossRefGoogle Scholar
  55. Linares OJ (2004) Bioestratigrafía de la fauna de mamíferos de las formaciones Socorro, Urumaco y Codore (Mioceno medio-Plioceno temprano) de la región de Urumaco, Falcón, Venezuela. Paleobiol Neotrop 1:1–26Google Scholar
  56. Lydekker R (1894) Contributions to the knowledge of the fossil vertebrates of Argentina. Part II. Cetacean skulls from Patagonia. An Mus La Plata 2:1–15Google Scholar
  57. Marx FG, Fordyce RE (2015) Baleen boom and bust: a synthesis of mysticete phylogeny, diversity and disparity. R Soc Open Sci 2. doi:  10.1098/rsos.140434
  58. Marx FG, Tsai C-H, Fordyce RE (2015) A new early Oligocene toothed ‘baleen’ whale (Mysticeti: Aetiocetidae) from western North America: one of the oldest and the smallest. R Soc Open Sci 2: 10.1098/rsos.150476
  59. Mead JG, Fordyce RE (2009) The therian skull: a lexicon with emphasis on the odontocetes. Smithson Contrib Zool 627:1–248CrossRefGoogle Scholar
  60. Mones A (1986) Palaeovertebrata Sudamericana. Catálogo sistemático de los vertebrados fósiles de America del Sur, Parte I. Lista preliminar y bibliografía. Courier Forschungsinstitut Senckenberg 82:1–625Google Scholar
  61. Moreno FP (1892) Noticias sobre algunos cetáceos fósiles y actuales de la República Argentina. Rev Mus La Plata 3:381–400Google Scholar
  62. Moreno F, Hendy AJW, Quiroz L, Hoyos N, Jones DS, Zapata V, Zapata S, Ballen GA, Cadena E, Cárdenas AL, Carrillo-Briceño JD, Carrillo JD, Delgado-Sierra D, Escobar J, Martínez JI, Martínez C, Montes C, Moreno J, Pérez N, Sánchez R, Suárez C, Vallejo-Pareja MC, Jaramillo C (2015) Revised stratigraphy of Neogene strata in the Cocinetas Basin, La Guajira, Colombia. Swiss J Palaeontol 134:5–43. doi:  10.1007/s13358-015-0071-4 CrossRefGoogle Scholar
  63. Muizon C de (1984) Les vertébrés fossiles de la Formation Pisco (Pérou). Deuxième partie: les odontocètes (Cetacea, Mammalia) du Pliocène inférieur de Sud-Sacaco. Trav Inst Fr E And 50:1–188Google Scholar
  64. Muizon C de (1988a) Les vertébrés fossiles de la Formation Pisco (Pérou). Troisème partie: Les odontocètes (Cetacea, Mammalia) du Miocène. Mem Inst Fr Et And 78:1–244Google Scholar
  65. Muizon C de (1988b) Le polyphylétisme des Acrodelphidae, odontocètes longisrostres du Miocène européen. Bull Mus Natl Hist Nat 10:31–88Google Scholar
  66. Muizon C de (1988c) Les relations phylogénétiques des Delphinida (Cetacea, Mammalia). Ann Paleontol 74:159–227Google Scholar
  67. Nummela S, Thewissen JGM, Bajpai S, Hussain T, Kumar K (2007) Sound transmission in archaic and modern whales: anatomical adaptations for underwater hearing. Anat Rec 290:716–733. doi:  10.1002/ar.20528 CrossRefGoogle Scholar
  68. O’Leary MA (2004) A fragmentary odontocete cranium from the lower Miocene of Venezuela. Spec Pap Palaeontol 71:99–104Google Scholar
  69. Ochsenius C (1980) Cuaternario en Venezuela. UNEFM, Coro, VenezuelaGoogle Scholar
  70. Odreman-Rivas OE (1997) Lista actualizada de los fósiles de Venezuela. In: La Marca E (ed) Vertebrados actuales y fósiles de Venezuela. Museo de ciencia y tecnología de Mérida, Mérida, Venezuela, pp 231–234Google Scholar
  71. Odreman-Rivas OE, Medina CJ (1984) Vertebrados fósiles de Venezuela secuencia, relaciones con otros países de América del Sur. Cuad Geol 1:60–86Google Scholar
  72. Omura H (1972) An osteological study of the Cuvier’s beaked whale, Ziphius cavirostris, in the Northwest Pacific. Sci Rep Whales Res Inst 24:1–34Google Scholar
  73. Pérez LM, Pérez Panera JP, Aguilera OA, Ronchi DI, Sánchez R, Manceñido MO, Sánchez-Villagra MR (2016) Palaeontology, sedimentology, and biostratigraphy of a fossiliferous outcrop of the early Miocene Querales Formation, Falcón Basin, Venezuela. Swiss J Palaeontol 135:187–203. doi:  10.1007/s13358-015-0105-y CrossRefGoogle Scholar
  74. Perrin WF (1975) Variation of spotted and spinner porpoise (genus Stenella) in the Eastern Pacific and Hawaii. Bull Scripps Inst Oceanogr 21:1–206Google Scholar
  75. Pilleri G, Gihr M (1979) Skull, sonar field and swimming behaviour or Ischyorhynchus vanbenedeni (Ameghino, 1891) and taxonomical position of the genera Ischyorhynchus, Anisodelphis and Pontoplanodes (Cetacea). Investig Cetacea 10:17–70Google Scholar
  76. Pyenson ND, Gutstein CS, Parham JF, Le Roux JP, Chavarría CC, Little H, Metallo A, Rossi V, Valenzuela-Toro AM, Velez-Juarbe J, Santelli CM, Rogers DR, Cozzuol MA, Suárez ME (2014) Repeated mass strandings of Miocene marine mammals from Atacama Region of Chile point to sudden death at sea. Proc R Soc Biol Sci Ser B 281:doi:  10.1098/rspb.2013.3316
  77. Pyenson ND, Vélez-Juarbe J, Gutstein CS, Little H, Vigil D, O’Dea A (2015) Isthminia panamensis, a new fossil inioid (Mammalia, Cetacea) from the Chagres Formation of Panama and the evolution of ‘river dolphins’ in the Americas. PeerJ 3:e1227. doi:  10.7717/peerj.1227 CrossRefPubMedPubMedCentralGoogle Scholar
  78. Quiroz L, Jaramillo C (2010) Stratigraphy and sedimentary environments of Miocene shallow to marginal marine deposits in the Urumaco Through, Falcón Basin, western Venezuela. In: Sánchez-Villagra MR, Aguilera O, Carlini AA (eds) Urumaco and Venezuelan Paleontology: The Fossil Record of the Northern Neotropics. Indiana University Press, Bloomington, pp 153–172Google Scholar
  79. Rey O (1994) Miembro Chiguaje: Formación La Vela, o Formación Codore. Boletín de la Sociedad Venezolana de Geólogos 19: 50–53Google Scholar
  80. Rey O (1996) Estratigrafía de la península de Paraguaná, Venezuela. Rev Fac Ing 11:36–45Google Scholar
  81. Rommel SA (1990) Osteology of the bottlenose dolphin. In: Leatherwood S, Reeves RR (eds) The Bottlenose Dolphin. Academic Press, San Diego, pp 29–49CrossRefGoogle Scholar
  82. Rommel SA, Reynolds JE, III (2008) Skeleton, postcranial. In: Perrin WF, Würsig B, Thewissen JGM (eds) Encyclopedia of Marine Mammals. Academic Press, San Diego, pp 1021–1032Google Scholar
  83. Sánchez‐Villagra MR, Aguilera OA (2006) Neogene vertebrates from Urumaco, Falcón State, Venezuela: diversity and significance. J Syst Palaeontol 4:213–220. doi:  10.1017/S1477201906001829 CrossRefGoogle Scholar
  84. Sánchez-Villagra MR, Aguilera OA, Sánchez R, Carlini AA (2010) The fossil vertebrate record of Venezuela of the last 65 million years. In: Sánchez-Villagra MR, Aguilera O, Carlini AA (eds) Urumaco and Venezuelan Paleontology: The Fossil Record of the Northern Neotropics. Indiana University Press, Bloomington, pp 19–51Google Scholar
  85. Sánchez-Villagra MR, Burnham RJ, Campbell DC, Feldmann RM, Gaffney ES, Kay RF, Lozsán R, Purdy R, Thewissen JGM (2000) A new near-shore marine fauna and flora from the early Neogene of northwestern Venezuela. J Paleontol 74:957–968Google Scholar
  86. Sánchez-Villagra MR, Gasparini Z, Lozsán R, Moody JM, Uhen MD (2001) New discoveries of vertebrates from a near-shore marine fauna from the early Miocene of northwestern Venezuela. Paläontol Z 75:227–232. doi:  10.1007/BF02988016 CrossRefGoogle Scholar
  87. Scheyer TM, Aguilera OA, Delfino M, Fortier DC, Carlini AA, Sánchez R, Carrillo-Briceño JD, Quiroz L, Sánchez-Villagra MR (2013) Crocodylian diversity peak and extinction in the late Cenozoic of the northern Neotropics. Nat Commun 4:1907. doi:  10.1038/ncomms2940 CrossRefPubMedGoogle Scholar
  88. Slijper EJ (1936) Die Cetaceen. Vergleischend-Anatomisch und Systematisch. Ein Beitrag zur vergleichenden Anatomie des Blutgefäss-, Nerven- und Muskelsystems, sowie des Rumpfskelettes der Säugetiere, mit Studien über die Theorie des Aussterbens und der Foetalisation. A. Asher & Co. B. V., AmsterdamGoogle Scholar
  89. Smith JB, Dodson P (2003) A proposal for a standard terminology of anatomical notation and orientation in fossil vertebrate dentitions. J Vertebr Paleontol 23:1–12CrossRefGoogle Scholar
  90. Smith CJ, Collins LS, Jaramillo C, Quiroz LI (2010) Marine paleonvironments of Miocene–Pliocene formations of north-central Falcón State, Venezuela. J Foraminifer Res 40:266–282. doi:  10.2113/gsjfr.40.3.266 CrossRefGoogle Scholar
  91. True FW (1910) Description of a skull and some vertebrae of the fossil cetacean Diochotichus vanbenedeni from Santa Cruz, Patagonia. Bull Am Mus Nat Hist 28: 19–32Google Scholar
  92. Uhen MD (2010) The origin(s) of whales. Annu Rev Earth Planet Sci 38:189–219. doi:  10.1146/annurev-earth-040809-152453 CrossRefGoogle Scholar
  93. Uhen MD, Coates AG, Jaramillo CA, Montes C, Pimiento C, Rincon A, Strong N, Velez-Juarbe J (2010) Marine mammals from the Miocene of Panama. J South Am Earth Sci 30:167–175. doi:  10.1016/j.jsames.2010.08.002 CrossRefGoogle Scholar
  94. Valerio AL, Laurito CA (2012) Cetáceos fósiles (Mammalia, Odontoceti, Eurhinodelphinoidea, Inioidea, Physeterioidea) de la Formación Curré, Mioceno superior (Hemphilliano temprano tardío) de Costa Rica. Rev Geol Am Cent 46:151–160Google Scholar
  95. Vélez-Juarbe J, Wood AR, De Gracia C, Hendy AJW (2015) Evolutionary patterns among living and fossil kogiid sperm whales: evidence from the Neogene of Central America. PLoS ONE 10:e0123909. doi:  10.1371/journal.pone.0123909 CrossRefPubMedPubMedCentralGoogle Scholar

Copyright information

© Springer Science+Business Media New York 2016

Authors and Affiliations

  1. 1.Paleontological Institute and MuseumUniversity of ZurichZürichSwitzerland
  2. 2.Center for Tropical Paleoecology and ArchaeologySmithsonian Tropical Research InstituteAncónPanama
  3. 3.Museo Paleontológico de UrumacoUrumacoVenezuela

Personalised recommendations