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 Volume 2 Chapter 6

Chapter  6

140 Million Years of Tropical Biome Evolution   


Citation is suggested as: 

Jaramillo, C. 2019. 140 million years of tropical biome evolution. In: Gómez, J. & Pinilla–Pachon, A.O. (editors), The Geology of Colombia, Volume 2 Mesozoic. Servicio Geológico Colombiano, Publicaciones Geológicas Especiales 36, 28 p. Bogotá.

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The origin and development of Neotropical biomes are central to our understanding of extant ecosystems and our ability to predict their future. During the Cretaceous, biomass of tropical rainforests was mostly dominated by gymnosperms and ferns, forest structure was poorly stratified and the canopy was open and dominated by gymnosperms. Extant tropical rainforests first developed at the onset of the Cenozoic, as a result of the massive extinction of the Cretaceous – Paleocene boundary. Paleocene rainforests were multistratified, with an angiosperm–dominated canopy that had high photosynthetic potential. Tropical climate has followed global patterns of warmings and coolings during the last 60 Ma. Rainforest diversity has increased during the warmings while it has decreased during coolings. Several extant biomes, including páramos, cloud forest, savannas, and dry/xerophytic forest, have increase significantly during the late Neogene at the expense of the reduction of the rainforest. Timing and drivers of these changes are still unknown but seem to be related to the onset of our modern, cool–state climate since the onset of the Pleistocene, 2.6 Ma ago.

Keywords:   Neotropical biomes, tropical rainforest, gymnosperms, angiosperms, evolution.


El origen y el desarrollo de los biomas neotropicales son fundamentales para nuestra comprensión de los ecosistemas actuales y nuestra capacidad para predecir su futuro. Durante el Cretácico, la biomasa de los bosques tropicales estaba dominada principalmente por gimnospermas y helechos, la estructura del bosque no poseía una estratificación marcada y el dosel era abierto y dominado por gimnospermas. Los bosques tropicales actuales se desarrollaron por primera vez al inicio del Cenozoico, como resultado de la extinción masiva del límite Cretácico–Paleoceno. Los bosques tropicales del Paleoceno eran multiestratificados, con un dosel dominado por angiospermas con alto potencial fotosintético. El clima tropical ha seguido patrones globales de calentamiento y enfriamiento durante los últimos 60 Ma. La diversidad del bosque tropical ha aumentado durante los calentamientos y disminuido durante los enfriamientos. Varios biomas que hoy existen, incluyendo páramos, bosques nubosos, sabanas y bosques secos/xerofíticos, han crecido significativamente desde el Neógeno tardío en áreas ocupadas previamente por el bosque tropical. Las causas y temporalidad de este cambio masivo en el paisaje aún se desconocen, pero parecen estar relacionadas con el inicio de nuestro clima frío moderno desde el comienzo del Pleistoceno, hace 2,6 Ma.

Palabras clave:  biomas neotropicales, bosque tropical, gimnospermas, angiospermas, evolución. 


PETM                 Paleocene Eocene Thermal Maximum

ETM                      Eocene Thermal Maximum

MMCO              Middle Miocene climatic optimum

GABI                   Great American Biotic Interchange

ITCZ                    Intertropical convergence zone

CAM                   Crassulacean acid metabolism

WUE                    Water use efficiency

DNA                    Deoxyribonucleic acid

WWF                  World Wildlife Fund


Aber, J., Neilson, R., McNulty, S., Lenihan, J.M., Bachelet, D. & Draper, R.J. 2001. Forest processes and global environmental change: Predicting the effects of individual and multiple stressors. BioScience, 51(9): 735–751.

Aguilera, O. 2004. Tesoros paleontológicos de Venezuela, Urumaco, patrimonio natural de la humanidad. Editorial Arte, 148 p. Caracas.

Aguilera, O., Lundberg, J., Birindelli, J., Sabaj–Pérez, M., Jaramillo, C.A. & Sánchez–Villagra, M.R. 2013a. Palaeontological evidence for the last temporal occurrence of the ancient western Amazonian River outflow into the Caribbean. PLOS ONE, 8(9): 1–17.

Aguilera, O., Moraes–Santos, H., Costa, S., Ohe, F., Jaramillo, C.A. & Nogueira, A. 2013b. 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 Journal of Palaeontology, 132(1): 45–68.

Aguilera, O., Andrade, G.O., Lopes, R.T., Machado, A.S., Dos Santos, T.M., Marques, G., Bertucci, T., Aguiar, T., Carrillo–Briceño, J., Rodríguez, F. & Jaramillo, C.A. 2017. Neogene proto–Caribbean porcupinefishes (Diodontidae). PLOS ONE, 12(7): 1–26.

Alda, F., Reina, R.G., Doadrio, I. & Bermingham, E. 2013. Phylogeny and biogeography of the Poecilia sphenops species complex (Actinopterygii, Poeciliidae) in Central America. Molecular Phylogenetics and Evolution, 66: 1011–1026.

Amson, E., Carrillo, J.D. & Jaramillo, C.A. 2016. Neogene sloth assemblages (Mammalia, Pilosa) of the Cocinetas Basin (La Guajira, Colombia): Implications for the Great American Biotic Interchange. Palaeontology, 59(4): 563–582.

Anderson, J.B., Warny, S., Askin, R.A., Wellner, J.S., Bohaty, S.M., Kirshner, A.E., Livsey, D.N., Simms, A.R., Smith, T.R., Ehrmann, W., Lawver, L.A., Barbeau, D., Wise, S.W., Kulhenek, D.K., Weaver, F.M. & Majewski, W. 2011. Progressive Cenozoic cooling and the demise of Antarctica's last refugium. Proceedings of the National Academy of Sciences, 108(28): 11356–11360.

Antoine, P.O., Salas–Gismondi, R., Baby, P., Benammi, M., Brusset, S., Franceshi, D., Espurt, N., Goillot, C., Pujos, F., Tejada, J. & Urbina, M. 2007. The middle Miocene (Laventan) Fitzcarrald fauna, Amazonian Peru. Cuadernos del Museo Geominero, 8: 19–24. Madrid.

Arakaki, M., Christin, P.A., Nyffeler, R., Lendel, A., Eggli, U., Ogburn, R.M., Spriggs, E., Moore, M.J. & Edwards, E.J. 2011. Contemporaneous and recent radiations of the world's major succulent plant lineages. Proceedings of the National Academy of Sciences, 108(20): 8379–8384.

Archangelsky, S. & Taylor, T.N. 1993. The ultrastructure of in situ Clavatipollenites pollen from the Early Cretaceous of Patagonia. American Journal of Botany, 80(8): 879–885.

Archer, D., Eby, M., Brovkin, V., Ridgwell, A., Cao, L., Mikolajewicz, U., Caldeira, K., Matsumoto, K., Munhoven, G., Montenegro, A. & Tokos, K. 2009. Atmospheric lifetime of fossil fuel carbon dioxide. Annual Review of Earth and Planetary Sciences, 37: 117–134 .

Bacon, C.D., Silvestro, D., Jaramillo, C., Smith, B.T., Chakrabarty, P. & Antonelli, A. 2015a. Biological evidence supports an early and complex emergence of the Isthmus of Panama. Proceedings of the National Academy of Sciences of the Unites States of America, 112(19): 6110–6115.

Bacon, C.D., Silvestro, D., Jaramillo, C., Smith, B.T., Chakrabarty, P. & Antonelli, A. 2015b. Reply to Lessios and Marko et al.: Early and progressive migration across the Isthmus of Panama is robust to missing data and biases. Proceedings of the National Academy of Sciences of the Unites States of America, 112(43): E5767–E5768.

Bacon, C.D., Molnar, P., Antonelli, A., Crawford, A.J., Montes, C. & Vallejo–Pareja, M.C. 2016. Quaternary glaciation and the Great American Biotic Interchange. Geology, 44(5): 375–378.

Bassow, S.L., McConnaughay, K.D. & Bazzaz, F.A. 1994. The response of temperate tree seedlings grown in elevated CO2 to extreme temperature events. Ecological Applications, 4(3): 593–603.

Bayona, G., Montes, C., Cardona, A., Jaramillo, C.A., Ojeda, G., Valencia, V. & Ayala–Calvo, C. 2011. Intraplate subsidence and basin filling adjacent to an oceanic arc–continent collision: A case from the southern Caribbean–South America plate margin. Basin Research, 23(4): 403–422.

Beaulieu, J.M., O'Meara, B.C., Crane, P.R. & Donoghue, M.J. 2015. Heterogeneous rates of molecular evolution and diversification could explain the Triassic age estimate for angiosperms. Systematic Biology, 64(5): 869–878.

Bell, C.D., Soltis, D.E. & Soltis, P.S. 2010. The age and diversification of the angiosperms re–revisited. American Journal of Botany, 97(8): 1296–1303.

Berry, J. & Björkman, O. 1980. Photosynthetic response and adaptation to temperature in higher plants. Annual Review of Plant Physiology, 31: 491–543.

Bice, K.L., Birgel, D., Meyers, P.A., Dahl, K.A., Hinrichs, K. & Norris, R.D. 2006. A multiple proxy and model study of Cretaceous upper ocean temperatures and atmospheric CO2 concentrations. Paleoceanography and Paleoclimatology, 21(2): 1–17.

Billups, K., Ravelo, A.C., Zachos, J.C. & Norris, R.D. 1999. Link between oceanic heat transport, thermohaline circulation, and the intertropical convergence zone in the early Pliocene Atlantic. Geology, 27(4): 319–322.<0319:LBOHTT>2.3.CO;2

Bloch, J.I., Woodruff, E.D., Wood, A.R., Rincón, A.F., Harrington, A.R., Morgan, G.S., Foster, D.A., Montes, C., Jaramillo, C., Jud, N.A., Jones, D.S. & MacFadden, B.J. 2016. First North American fossil monkey and early Miocene tropical biotic interchange. Nature, 533(7602): 243–246.

Bowen, G.J. & Zachos, J.C. 2010. Rapid carbon sequestration at the termination of the Palaeocene – Eocene Thermal Maximum. Nature Geoscience, 3: 866–869.

Boyce, C.K. & Lee, J.E. 2010. An exceptional role for flowering plant physiology in the expansion of tropical rainforests and biodiversity. Proceedings of the Royal Society of London Series B: Biological Sciences, 277: 3437–3443.

Bralower, T., Thomas, D.J., Thomas, E. & Zachos, J.C. 1997. High–resolution records of the late Paleocene Thermal Maximum and circum–Caribbean volcanism: Is there a causal link? Geology, 25(11): 963–966.<0963:HRROTL>2.3.CO;2

Brenner, G. 1974. Palynostratigraphy of the Lower Cretaceous Gevar'am and Talme Yafe Formations in the Gever'am 2 well (southern coastal plain Israel). Geological Survey of Israel Bulletin, 59: 1–27.

Burnham, R.J. 2009. An overview of the fossil record of climbers: Bejucos, sogas, trepadoras, lianas, cipós, and vines. Revistra Brasileira de Paleontologia, 12(2): 149–160.

Burnham, R.J. & Graham, A. 1999. The history of Neotropical vegetation: New developments and status. Annals of the Missouri Botanical Garden, 86(2): 546–589.

Burnham, R.J. & Johnson, K.R. 2004. South American palaeobotany and the origins of Neotropical rainforests. Philosophical Transactions of the Royal Society of London Series B: Biological Sciences, 359(1450): 1595–1610.

Cadena, E.A. & Jaramillo, C. 2006. New Podocnemididae fossil turtles from the late Paleocene Cerrejón Formation, Guajira Peninsula, Colombia. 66th Annual Meeting of the Society of Vertebrate Paleontology. Unpublished Field Trip Guidebook, 43 p. Ottawa, Canada.

Cadena, E.A. & Jaramillo, C. 2015. The first fossil skull of Chelus (Pleurodira: Chelidae, Matamata turtle) from the early Miocene of Colombia. Palaeontologia Electronica, (18.2.32A): 1–10.

Cadena, E.A. & Schweitzer, M.H. 2014. A pelomedusoid turtle from the Paleocene – Eocene of Colombia exhibiting preservation of blood vessels and osteocytes. Journal of Herpetology, 48(4): 461–465.

Cadena, E.A., Bloch, J. & Jaramillo, C. 2012a. New bothremydid turtle (Testudines, Pleurodira) from the Paleocene of northeastern Colombia. Journal of Paleontology, 86(4): 688–698.

Cadena, E.A., Bourque, J., Rincón, A., Bloch, J.I., Jaramillo, C. & MacFadden, B. 2012b. New turtles (Chelonia) from the late Eocene through late Miocene of the Panama Canal Basin. Journal of Paleontology, 86(3): 539–557.

Cadena, E.A., Ksepka, D.T., Jaramillo, C. & Bloch, J.I. 2012c. New pelomedusoid turtles from the late Palaeocene Cerrejón Formation of Colombia and their implications for phylogeny and body size evolution. Journal of Systematic Palaeontology, 10(2): 313–331.

Carrillo, J.D., Forasiepi, A., Jaramillo, C. & Sánchez–Villagra, M.R. 2015. Neotropical mammal diversity and the Great American Biotic Interchange: Spatial and temporal variation in South America's fossil record. Frontiers in Genetics, 5(451): 1–12.

Boyce, C.K. & Lee, J.E. 2010. An exceptional role for flowering plant physiology in the expansion of tropical rainforests and biodiversity. Proceedings of the Royal Society of London Series B: Biological Sciences, 277: 3437–3443.

Carvalho, M.R., Herrera, F., Jaramillo, C.A., Wing, S. & Callejas, R. 2011. Paleocene Malvaceae from northern South America and their biogeographical implications. American Journal of Botany, 98(8): 1337–1355.

Carvalho, M.R., Wilf, P., Barrios, H., Windsor, D.M., Currano, E., Labandeira, C. & Jaramillo, C. 2014. Insect leaf–chewing damage tracks herbivore richness in modern and ancient forests. PLOS ONE, 9(5): 1–9.

Cernusak, L.A., Winter, K., Martínez, C., Correa, E., Aranda, J., García, M., Jaramillo, C. & Turner, B.L. 2011. Responses of legume versus nonlegume tropical tree seedlings to elevated CO2 concentration. Plant Physiology, 157: 372–385.

Cernusak, L.A., Winter, K., Dalling, J.W., Holtum, J.A.M., Jaramillo, C., Körner, C., Leakey, A.D.B., Norby, R.J., Poulter, B., Turner, B.L. & Wright, S.J. 2013. Tropical forest responses to increasing atmospheric CO2: Current knowledge and opportunities for future research. Functional Plant Biology, 40(6): 531–551.

Chaisson, W.P. 1995. Planktonic foraminiferal assemblages and paleoceanographic change in the trans–tropical Pacific Ocean: A comparison of west (Leg 130) and east (Leg 138), latest Miocene to Pleistocene. In: Pisias, N.G., Mayer, L.A., Jenecek, T.R., Palmer–Julson, A. & van Andel, T.H. (editors), Proceedings of the Ocean Drilling Program, Scientific Results 138, p. 555–597.

Chaisson, W.P. & Ravelo, A.C. 1997. Changes in upper water–column structure at Site 925, late Miocene – Pleistocene: Planktonic foraminifer assemblage and isotopic evidence. In: Shackleton, N.J., Curry, W.B., Richter, C. & Bralower, T.J. (editors), Proceedings of the Ocean Drilling Program, Scientific Results 154, p. 255–268.

Chiang, J.C.H. 2009. The tropics in paleoclimate. Annual Review of Earth and Planetary Sciences, 37: 263–297.

Chiang, J.C.H. & Bitz, C.M. 2005. Influence of high latitude ice cover on the marine intertropical convergence zone. Climate Dynamics, 25(5): 477–496.

Clyde, W.C. & Gingerich, P.D. 1998. Mammalian community response to the latest Paleocene Thermal Maximum: An isotaphonomic study in the northern Bighorn Basin, Wyoming. Geology, 26(11): 1011–1014.<1011:MCRTTL>2.3.CO;2

Cody, S., Richardson, J.E., Rull, V., Ellis, C. & Pennington, R.T. 2010. The Great American Biotic Interchange revisited. Ecography, 33(2): 326–332.

Connell, J.H. 1971. On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees. In: den Boer, P.J. & Gradwell, G.R. (editors), Dynamics of populations. Centre for Agricultural Publication and Documentation, p. 298–312. Wageningen, the Netherlands.

Correa, E., Jaramillo, C., Manchester, S. & Gutiérrez, M. 2010. A fruit and leaves of rhamnaceous affinities from the Late Cretaceous (Maastrichtian) of Colombia. American Journal of Botany, 97(1): 71–79.

Cozzuol, M. 2006. The Acre vertebrate fauna: Age, diversity, and geography. Journal of South American Earth Sciences, 21(3): 185–203.

Crane, P.R. & Lidgard, S. 1989. Angiosperm diversification and paleolatitudinal gradients in Cretaceous floristic diversity. Science, 246(4930): 675–678.

Crane, P.R. & Lidgard, S. 1990. Angiosperm radiation and patterns of Cretaceous palynological diversity. In: Taylor, P.D. & Larwood, G.P. (editors), Major evolutionary radiations 42, p. 377–407. Oxford.

Crifò, C., Currano, E.D., Baresh, A. & Jaramillo, C. 2014. Variations in angiosperm leaf vein density have implications for interpreting life form in the fossil record. Geology, 42(10): 919–922.

Currano, E.D., Wilf, P., Wing, S.L., Labandeira, C.C., Lovelock, E.C. & Royer, D.L. 2008. Sharply increased insect herbivory during the Paleocene – Eocene Thermal Maximum. Proceedings of the National Academy of Sciences of the United States of America, 105(6): 1960–1964.

Davis, C., Webb, C.O., Wurdack, K.J., Jaramillo, C. & Donoghue, M.J. 2005. Explosive radiation of Malpighiales supports a mid–Cretaceous origin of modern tropical rain forests. The American Naturalist, 165(3): E36–E65.

De Boer, B., van de Wal, R.S.W., Bintanja, R., Lourens, L.J. & Tuenter, E. 2010. Cenozoic global ice–volume and temperature simulations with 1–D ice–sheet models forced by benthic δ18O records. Annals of Glaciology, 51(55): 23–33.

De Boer, H.J., Lammertsma, E.I., Wagner–Cremer, F., Dilcher, D.L., Wassen, M.J. & Dekker, S.C. 2011. Climate forcing due to optimization of maximal leaf conductance in subtropical vegetation under rising CO2. Proceedings of the National Academy of Sciences of the United States of America, 108(10): 4041–4046.

De la Parra, F. 2009. Palynological changes across the Cretaceous – Tertiary boundary in Colombia, South America. Master thesis, University of Florida, 105 p. Gainesville, USA.

De la Parra, F., Jaramillo, C. & Dilcher, D. 2008a. Paleoecological changes of spore producing plants through the Cretaceous – Paleocene boundary in Colombia. Palynology, 32: 258–259.

De la Parra, F., Jaramillo, C., Rueda, M. & Dilcher, D. 2008b. Has there been a plant mass extinction in the last 70 million years in the Neotropics? 12th International Palynological Congress. Proceedings, p. 59. Bonn, Germany.

Díaz de Gamero, M.L. & Linares, O.J. 1989. Estratigrafía y paleontología de la Formación Urumaco, del Mioceno tardío de Falcón noroccidental. 7th Congreso Geológico Venezolano. Proceedings, 1, p. 419–439. Caracas.

Dickens, G.R., O'Neil, J.R., Rea, D.K. & Owen, R.M. 1995. Dissociation of oceanic methane hydrate as a cause of the carbon isotope excursion at the end of the Paleocene. Paleoceanography and Paleoclimatology, 10(6): 965–971.

Dickens, G.R., Bralower, T.J., Thomas, D.J., Thomas, E. & Zachos, J.C. 1998. High–resolution records of the late Paleocene Thermal Maximum and circum–Caribbean volcanism: Is there a causal link?: Comment and reply. Geology, 26(7): 670–671.<0670:HRROTL>2.3.CO;2

Doria, G., Jaramillo, C. & Herrera, F. 2008. Menispermaceae from the Cerrejón Formation, middle to late Paleocene, Colombia. American Journal of Botany, 95(8): 954–973.

Doubinger, J. 1973. Pollen and spores from the Paleocene coal basin of Cerrejon (Guajira Province, Colombia). Comptes Rendus du 96 Congres National des Societes Savantes. Proceedings, 5, p. 253–262. Tolouse, France.

Doyle, J.A. 2012. Molecular and fossil evidence on the origin of angiosperms. Annual Review of Earth and Planetary Science, 40: 301–326.

Doyle, J.A. & Hickey, L.J. 1976. Pollen and leaves from the mid–Cretaceous Potomac Group and their bearing on early angiosperm evolution. In: Beck, C.B. (editor), Origin and early evolution of angiosperms. Columbia University Press, p. 139–206. New York.

Doyle, J.A., Biene, P., Doerenkamp, A. & Jardine, S. 1977. Angiosperm pollen from the pre–Albian Lower Cretaceous of Equatorial Africa. Bulletin des Centres de Recherches Exploration–Production Elf–Aquitaine, 1: 451–473.

Doyle, J.A., Hotton, C. & Ward, J. 1990. Early Cretaceous tetrads, zonasulculate pollen, and Winteraceae. I. Taxonomy, morphology, and ultrastructure. American Journal of Botany, 77(12): 1544–1557.

Edwards, E.J., Osborne, C.P., Stromberg, C.A.E., Smith, S.A., Bond, W.J., Christin, P.A., Cousins, A.B., Duvall, M.R., Fox, D.L., Freckleton, R.P., Ghannoum, O., Hartwell, J., Huang, Y., Janis, C.M., Keeley, J.E., Kellogg, E.A., Knapp, A.K., Leakey, A.D.B., Nelson, D.M., Saarela, J.M., Sage, R.F., Sala, O.E., Salamin, N., Still, C.J. & Tipple, B. 2010. The origins of C4 grasslands: Integrating evolutionary and ecosystem science. Science, 328(5978): 587–591.

Elmer, K.R., Bonett, R.M., Wake, D.B. & Lougheed, S. 2013. Early Miocene origin and cryptic diversification of South American salamanders. BMC Evolutionary Biology, 13(59): 1–16.

Erwin, D.H. 2008. Extinction: How life on earth nearly ended 250 million years ago. Princeton University Press, 320 p. Princeton.

Farris, D.W., Jaramillo, C., Bayona, G., Restrepo–Moreno, S.A., Montes, C., Cardona, A., Mora, A., Speakman, R.J., Glascock, M.D. & Valencia, V. 2011. Fracturing of the Panamanian Isthmus during initial collision with South America. Geology, 39(11): 1007–1010.

Fedorov, A.V., Brierley, C.M., Lawrence, K.T., Liu, Z., Dekens, P.S. & Ravelo, A.C. 2013. Patterns and mechanisms of early Pliocene warmth. Nature, 496(7443): 43–49.

Feild, T.S., Brodribb, T.J., Iglesias, A., Chatelet, D.S., Baresh, A., Upchurch, G.R., Gómez, B., Mohr, B.A.R., Coiffard, C., Kvaček, J. & Jaramillo, C.A. 2011a. Fossil evidence for Cretaceous escalation in angiosperm leaf vein evolution. Proceedings of the National Academy of Sciences of the United States of America, 108(20): 8363–8366.

Feild, T.S., Upchurch, G.R., Chatelet, D.S., Brodribb, T.J., Grubbs, K.C., Samain, M.S. & Wanke, S. 2011b. Fossil evidence for low gas exchange capacities for Early Cretaceous angiosperm leaves. Paleobiology, 37(2): 195–213.

Figueiredo, J., Hoorn, C., van der Ven, P. & Soares, E. 2009. Late Miocene onset of the Amazon River and the Amazon deep–sea fan: Evidence from the Foz do Amazonas Basin. Geology, 37(7): 619–622.

Filippelli, G.M. & Flores, J.A. 2009. From the warm Pliocene to the cold Pleistocene: A tale of two oceans. Geology, 37(10): 959–960.

Fine, P.V.A. & Ree, R.H. 2006. Evidence for a time–integrated species–area effect on the latitudinal gradient in tree diversity. The American Naturalist, 168(6): 796–804.

Fine, P.V.A., Ree, R.H. & Burnham, R.J. 2008. Disparity in tree species richness between tropical, temperate and boreal biomes. The geographic area and age hypothesis. In: Carson, W.P. & Schnitzer, S.A. (editors), Tropical forest community ecology. Blackwell Scientific, p. 31–45. London.

Flohn, H. 1981. A hemispheric circulation asymmetry during late Tertiary. Geologische Rundschau, 70: 725–736.

Forasiepi, A.M., Soibelzon, L.H., Suárez, C., Sánchez, R., Quiroz, L.I., Jaramillo, C. & Sánchez–Villagra, M.R. 2014. Carnivorans at the Great American Biotic Interchange: New discoveries from the northern Neotropics. Naturwissenschaften, 101(11): 965–974.

Frailey, C.D. 1986. Late Miocene and Holocene mammals, exclusive of the Notoungulata, of the Rio Acre region, western Amazonia. Contribution in Sciences, 374: 1–46.

Frieling, J., Gebhardt, A., Hubert, M., Adekeye, O.A., Okande, S.O., Reichart, G.J., Middelburg, J.J., Schouten, S. & Sluijs, A. 2017. Extreme warmth and heat–stressed plankton in the tropics during the Paleocene – Eocene Thermal Maximum. Science Advances, 3(3): 1–9.

Friis, E.M., Crane, P.R., Pedersen, K.R., Stampanoni, M. & Marone, F. 2015. Exceptional preservation of tiny embryos documents seed dormancy in early angiosperms. Nature, 528: 551–554.

García, C. 1958. Investigación palinológica de la Formación Guaduas del Anticlinal de Guachetá–Lenguazaque–Tausa. Boletín de Geología, (2): 27–31.

Garzione, C.N., Molnar, P., Libarkin, J.C. & MacFadden, B.J. 2006. Rapid late Miocene rise of the Bolivian altiplano: Evidence for removal of mantle lithosphere. Earth and Planetary Science Letters, 241(3–4): 543–556.

Garzione, C.N., Hoke, G.D., Libarkin, J.C., Withers, S., MacFadden, B., Eiler, J., Ghosh, P. & Mulch, A. 2008. Rise of the Andes. Science, 320(5881): 1304–1307.

Garzione, C.N., Auerbach, D.J., Smith, J.J.S., Rosario, J.J., Passey, B.H., Jordan, T.E. & Eiler, J.M. 2014. Clumped isotope evidence for diachronous surface cooling of the altiplano and pulsed surface uplift of the central Andes. Earth and Planetary Science Letters, 393: 173–181.

Gaston, K.J. 2000. Global patterns in biodiversity. Nature, 405: 220–227.

Gehler, A., Gingerich, P.D. & Pack, A. 2016. Temperature and atmospheric CO2 concentration estimates through the PETM using triple oxygen isotope analysis of mammalian bioapatite. Proceedings of the National Academy of Sciences of the United States of America, 113(28): 7739–7744.

Gentry, A.H. 1982a. Neotropical floristic diversity: Phytogeographical connections between Central and South America, Pleistocene climatic fluctuations, or an accident of the Andean Orogeny? Annals of the Missouri Botanical Garden, 69(3): 557–593.

Gentry, A.H. 1982b. Patterns of Neotropical plant species diversity. Evolutionary Biology, 15: 1–84.

Ghosh, P., Garzione, C.N. & Eiler, J.M. 2006. Rapid uplift of the altiplano revealed through 13C–18O bonds in paleosol carbonates. Science, 311(5760): 511–515.

Gillett, J.B. 1962. Pest pressure, an underestimated factor in evolution. Systematics Association Publication, 4: 37–46.

Gingerich, P.D. 2006. Environment and evolution through the Paleocene – Eocene Thermal Maximum. Trends in Ecology & Evolution, 21(5): 246–253.

González, H. 1977. Flora de la Formación Valle Alto, Jurásico, en la cordillera Central de Colombia. Boletín Ciencias de la Tierra, 2: 107–122.

Gómez–Navarro, C., Jaramillo, C., Herrera, F., Wing, S.L. & Callejas, R. 2009. Palms (Arecaceae) from a Paleocene rainforest of northern Colombia. American Journal of Botany, 96(7): 1300–1312.

González–Wevar, C.A., Hüne, M., Segovia, N.I., Nakano, T., Spencer, H.G., Chown, S.L., Saucede, T., Johnstone, G., Mansilla, A. & Poulin, E. 2016. Following the Antarctic Circumpolar Current: Patterns and processes in the biogeography of the limpet Nacella (Mollusca: Patellogastropoda) across the Southern Ocean. Journal of Biogeography, 44(4): 861–874.

Graham, A. 1988a. Studies in Neotropical paleobotany. VI. The lower Miocene communities of Panama–The Cucaracha Formation. Annals Missouri Botanical Garden, 75(4): 1467–1479.

Graham, A. 1988b. Studies in Neotropical paleobotany. V. The lower Miocene communities of Panama–The Culebra Formation. Annals Missouri Botanical Garden, 75(4): 1440–1466.

Graham, A. 1991. Studies in Neotropical paleobotany. X. The Pliocene communities of Panama–composition, numerical representations, and paleocommunity paleoenvironmental reconstructions. Annals Missouri Botanical Garden, 78(2): 465–475.

Graham, A. 1992. Utilization of the isthmian land bridge during the Cenozoic–paleobotanical evidence for timing, and the selective influence of altitudes and climate. Review of Palaeobotany and Palynology, 72(1–2): 119–128.

Graham, A. 1999. Late Cretaceous and Cenozoic history of North American vegetation. Oxford University Press, 350 p. New York.

Graham, A., editor. 2010. Late Cretaceous and Cenozoic history of Latin American vegetation and terrestrial environments. Missouri Botanical Garden Press, 618 p. Saint Louis, USA.

Graham, A. 2011. The age and diversification of terrestrial new world ecosystems through Cretaceous and Cenozoic time. American Journal of Botany, 98(3): 336–351.

Gübeli, A.A., Hochuli, P. & Wildi, W. 1984. Lower Cretaceous turbiditic sediments from the Rif chain (northern Marocco)–palynology, stratigraphy and palaeogeographic setting. Geologische Rundschau, 73(3): 1081–1114.

Gutiérrez, N.M. & Jaramillo, C.A. 2007. Maastrichtian paleotemperature and paleoprecipitation from the Guaduas Formation, Colombia. Palynology, 32: 260.

Gutjar, M., Ridgwell, A., Sexton, P.F., Anagnostou, E., Pearson, P.N., Pälike, H., Norris, R.D., Thomas, E. & Foster, G.L. 2017. Very large release of mostly volcanic carbon during the Palaeocene – Eocene Thermal Maximum. Nature, 548: 573–577.

Haffer, J. 1969. Speciation in Amazonian forest birds. Science, 165(3889): 131–137.

Hambalek, N. 1993. Palinoestratigrafia del Mioceno–Plioceno de la región de Urumaco, Falcon noroccidental. Bachelor thesis, Universidad Central de Venezuela, 168 p. Caracas.

Hambalek, N., Rull, V., Digiacomo, E. & Díaz de Gamero, M.L. 1994. Evolución paleoecológica y paleoambiental de la secuencia del Neógeno en el surco de Urumaco: Estudio palinológico y litológico. Boletín de la Sociedad Venezolana de Geología, 19: 7–19.

Haq, B.U., Hardenbol, J., Vail, P.R., Stover, L.E., Colin, J.P., Ioannides, N.S., Wright, R.C., Baum, G.R., Gombos–Jr, A.M., Pflum, C.E., Loutit, T.S., du Chêne, R.J., Romine, K.K., Sarg, J.F., Posamentier, H.W. & Morgan, B.E. 1988. Mesozoic and Cenozoic chronostratigraphy and cycles of sea–level change. In: Wilgus, C.K., Hastings, B.S., Posamentier, H., van Wagoner, J., Ross, C.A. & Kendall, C.G. (editors), Sea–level changes: An integrated approach. Society of Economic Paleontologists and Mineralogists, Special Publication 42, p. 71–108.

Hastings, A., Bloch, J., Cadena, E. & Jaramillo, C. 2010. A new small short–snouted dyrosaurid (Crocodylomorpha, Mesoeucrocodylia) from the Paleocene of northeastern Colombia. Journal of Vertebrate Paleontology, 30(1): 139–162.

Hastings, A., Bloch, J. & Jaramillo, C. 2011. A new longirostrine dyrosaurid (Crocodylomorpha, Mesoeucrocodylia) from the Paleocene of north–eastern Colombia: Biogeographic and behavioural implications for new–world Dyrosauridae. Palaeontology, 54(5): 1095–1116.

Hastings, A., Bloch, J., Jaramillo, C., Rincón, A. & MacFadden, B. 2013. Systematics and biogeography of crocodylians from the Miocene of Panama. Journal of Vertebrate Paleontology, 33(2): 239–263.

Hastings, A., Bloch, J. & Jaramillo, C. 2014. A new blunt–snouted dyrosaurid, Anthracosuchus balrogus gen. et sp. nov. (Crocodylomorpha, Mesoeucrocodylia), from the Palaeocene of Colombia. Historical Biology: An International Journal of Paleobiology, 27(8): 998–1020.

Haug, G. & Tiedemann, R. 1998. Effect of the formation of the Isthmus of Panama on Atlantic Ocean thermohaline circulation. Nature, 393: 673–676.

Haug, G.H., Hughen, K.A., Sigman, D.M., Peterson, L.C. & Röhl, U. 2001. Southward migration of the intertropical convergence zone through the Holocene. Science, 293(5533): 1304–1308.

Head, J., Bloch, J., Hasting, A., Bourque, J., Cadena, E., Herrera, F., Polly, P.D. & Jaramillo, C. 2009a. Giant boid snake from the Palaeocene Neotropics reveals hotter past equatorial temperatures. Nature, 457(7230): 715–718.

Head, J., Bloch, J., Hasting, A., Bourque, J., Cadena, E., Herrera, F., Polly, P.D. & Jaramillo, C. 2009b. Head et al. reply. Nature, 460: E4–E5.

Head, J., Rincón, A., Suárez, C., Montes, C. & Jaramillo, C. 2012. Fossil evidence for earliest Neogene American faunal interchange: Boa (Serpentes, Boinae) from the early Miocene of Panama. Journal of Vertebrate Paleontology, 32(6): 1328–1334.

Hendy, A.J.W., Jones, D.S., Moreno, F., Zapata, V. & Jaramillo, C. 2015. Neogene molluscs, shallow marine paleoenvironments, and chronostratigraphy of the Guajira Peninsula, Colombia. Swiss Journal of Palaeontology, 134(1): 45–75.

Herendeen, P.S., Friis, E.M., Pedersen, K.R. & Crane, P.R. 2017. Palaeobotanical redux: Revisiting the age of the angiosperms. Nature Plants, 3(17015): 1–8.

Herngreen, G.F.W. & Dueñas, H. 1990. Dating of the Cretaceous Une Formation, Colombia and the relationship with the Albian – Cenomanian African–South American microfloral province. Review of Palaeobotany and Palynology, 66(3–4): 345–359.

Herngreen, G.F.W., Kedves, M., Rovnina, L.V. & Smirnova, S.B. 1996. Cretaceous palynofloral provinces: A review. In: Jansonius, J. & McGregor, D.C. (editors), Palynology: Principles and applications. American Association of Stratigraphic Palynologists Foundation, 3, p. 1157–1188. Dallas.

Herrera, F., Jaramillo, C., Dilcher, D., Wing, S.L. & Gómez, C. 2008. Fossil Araceae from a Paleocene Neotropical rainforest in Colombia. American Journal of Botany, 95(12): 1569–1583.

Herrera, F., Manchester, S., Jaramillo, C., MacFadden, B. & da Silva–Caminha, S. 2010. Phytogeographic history and phylogeny of the Humiriaceae. International Journal of Plant Sciences, 171(4): 2004–2017

Herrera, F., Manchester, S.R., Hoot, S.B., Wefferling, K., Carvalho, M. & Jaramillo, C. 2011. Phytogeographic implications of fossil endocarps of Menispermaceae from the Paleocene of Colombia. American Journal of Botany, 98(12): 1–14.

Herrera, F., Manchester, S., Vélez–Juarbe, J. & Jaramillo, C.A. 2014a. Phytogeographic history of the Humiriaceae (Part 2). International Journal of Plant Science, 175(7): 828–840.

Herrera, F., Manchester, S.R., Carvalho, M.R., Jaramillo, C. & Wing, S.L. 2014b. Paleocene wind–dispersed fruits and seeds from Colombia and their implications for early Neotropical rainforests. Acta Palaeobotanica, 54(2): 197–229.

Herrera, F., Manchester, S.R., Koll, R. & Jaramillo, C. 2014c. Fruits of Oreomunnea (Juglandaceae) in the early Miocene of Panama. In: Stevens, W.D., Montiel, O.M. & Raven, P. (editors), Paleobotany and biogeography: A festschrift for Alan Graham in his 80th year. Missouri Botanical Garden Press, p. 124–133. Saint Louis, USA.

Hinojosa, F. & Villagran, C. 2005. Did South American mixed paleofloras evolve under thermal equability or in the absence of an effective Andean barrier during the Cenozoic? Palaeogeography, Palaeoclimatology, Palaeoecology, 217(1–2): 1–23.

Hooghiemstra, H. & van der Hammen, T. 1998. Neogene and Quaternary development of the Neotropical rain forest: The forest refugia hypothesis, and a literature overview. Earth–Science Reviews, 44(3–4): 147–183.

Hooghiemstra, H., Wijninga, V.M. & Cleef, A.M. 2006. The paleobotanical record of Colombia: Implications for biogeography and biodiversity. Annals of the Missouri Botanical Garden, 93(2): 297–325.

Hoorn, C. 1994a. An environmental reconstruction of the palaeo–Amazon River system (middle – late Miocene, NW Amazonia). Palaeogeography, Palaeoclimatology, Palaeoecology, 112(3–4): 187–238.

Hoorn, C. 1994b. Fluvial palaeonvironments in the intracratonic Amazonas Basin (early Miocene – early middle Miocene, Colombia). Palaeogeography, Palaeoclimatology, Palaeoecology, 109(1): 1–54.

Hoorn, C., Guerrero, J., Sarmiento, G.A. & Lorente, M.A. 1995. Andean tectonics as a cause for changing drainage patterns in Miocene northern South America. Geology, 23(3): 237–240.<0237:ATAACF>2.3.CO;2

Hoorn, C., Wesselingh, F.P., ter Steege, H., Bermúdez, M.A., Mora, A., Sevink, J., Sanmartín, I., Sánchez–Meseguer, A., Anderson, C.L., Figueiredo, J.P., Jaramillo, C., Riff, D., Negri, F.R., Hooghiemstra, H., Lundberg, J., Stadler, T., Sarkinen, T. & Antonelli, A. 2010. Amazonia through time: Andean uplift, climate change, landscape evolution, and biodiversity. Science, 330(6006): 927–931.

Hoorn, C., Bogotá, G.R., Romero–Báez, M., Lammertsma, E.I., Flantua, S.G.A., Dantas, E.L., Dino, R., do Carmo, D.A. & Chemale Jr., F. 2017. The Amazon at sea: Onset and stages of the Amazon River from a marine record, with special reference to Neogene plant turnover in the drainage basin. Global and Planetary Change, 153: 51–65.

Hotton, C.L. 2002. Palynology of the Cretaceous – Tertiary boundary in Central Montana: Evidence for extraterrestrial impact as a cause of the terminal Cretaceous extinctions. In: Hartman, J.H., Johnson, K.R. & Nichols, D.J. (editors), The Hell Creek Formation and the Cretaceous – Tertiary boundary in the northern great plains: An integrated continental record of the end of the Cretaceous. Geological Society of America, Special Paper 361, p. 473–502. Boulder, USA.

Hovan, S. 1995. Late Cenozoic atmospheric circulation intensity and climatic history recorded by eolian deposition in the eastern equatorial Pacific Ocean, Leg 138. In: Pisias, N.G., Mayer, L.A., Janecek, T.R., Palmer–Julson, A. & van Andel, T.H. (editors), Proceedings of the Ocean Drilling Program. Scientific Results 138, p. 615–625.

Huber, M. 2008. A hotter greenhouse? Science, 321(5887): 353–354.

Huber, M. 2009. Snakes tell a torrid tale. Nature, 457: 669–671.

Huber, M. & Caballero, R. 2011. The early Eocene equable climate problem revisited. Climate of the Past, 7: 603–633.

Huber, M. & Sloan, L.C. 1999. Warm climate transitions: A general circulation modeling study of the late Paleocene Thermal Maximum (~56 Ma). Journal of Geophysical Research: Atmospheres, 104(D14): 16633–16655.

Huber, M. & Sloan, L.C. 2000. Climatic responses to tropical sea surface temperature changes on a "greenhouse" Earth. Paleoceanography and Paleoclimatology, 15(4): 443–450.

Huber, M., Sloan, L.C. & Shellito, C. 2003. Early Paleogene oceans and climate: A fully coupled modeling approach using the NCAR CCSM. In: Wing, S.L., Gingerich, P.D., Schmitz, B. & Thomas, E. (editors), Causes and consequences of globally warm climates in the early Paleogene. Geological Society of America, Special Paper 369, p. 25–47. Boulder, USA.

Huertas, G. 2003. Flora fósil de Villa de Leyva y sus alrededores (Boyacá, Colombia, Suramérica). Camargo Editores, 151 p. Chía, Colombia.

Jablonski, D. 1993. The tropics as a source of evolutionary novelty through geological time. Nature, 364: 142–144.

Jacobs, B., Kingston, J. & Jacobs, L. 1999. The origin of grass–dominated ecosystems. Annals of the Missouri Botanical Garden, 86(2): 590–643.

Jacques, F.M., Wang, W., Ortiz, R., Li, H.L., Zhou, Z.K. & Chen, Z. 2011. Integrating fossils in a molecular–based phylogeny and testing them as calibration points for divergence time estimates in Menispermaceae. Journal of Systematics and Evolution, 49(1): 25–49.

Janzen, D.H. 1970. Herbivores and the number of tree species in tropical forests. The American Naturalist, 104(940): 501–528.

Jaramillo, C. 2002. Response of tropical vegetation to Paleogene warming. Paleobiology, 28(2): 222–243.<0222:ROTVTP>2.0.CO;2

Jaramillo, C. 2018. Evolution of the Isthmus of Panama: Biological, paleoceanographic, and paleoclimatological implications. In: Hoorn, C., Perrigo, A. & Antonelli, A. (editors), Mountains, climate and biodiversity. Wiley–Blackwell, p. 323–338. Chichester, UK.

Jaramillo, C. & Cárdenas, A. 2013. Global warming and Neotropical rainforests: A historical perspective. Annual Review of Earth and Planetary Sciences, 41: 741–766.

Jaramillo, C. & Dilcher, D.L. 2000. Microfloral diversity patterns of the late Paleocene – Eocene interval in Colombia, northern South America. Geology, 28(9): 815–818.<815:MDPOTL>2.0.CO;2

Jaramillo, C. & Dilcher, D.L. 2001. Middle Paleogene palynology of central Colombia, South America: A study of pollen and spores from tropical latitudes. Palaeontographica Abteilung B, 258(4–6): 87–213.

Jaramillo, C., Rueda, M. & Mora, G. 2006. Cenozoic plant diversity in the Neotropics. Science, 311(5769): 1893–1896.

Jaramillo, C., Bayona, G., Pardo–Trujillo, A., Rueda, M., Torres, V., Harrington, G. & Mora, G. 2007. The palynology of the Cerrejón Formation (upper Paleocene) of northern Colombia. Palynology, 31(1): 153–189.

Jaramillo, C., Hoorn, C., Silva, S., Leite, F., Herrera, F., Quiroz, L., Dino, R. & Antonioli, L. 2010a. The origin of the modern Amazon rainforest: Implications of the palynological and palaeobotanical record. In: Hoorn, C. & Wesselingh, F.P. (editors), Amazonia: Landscape and species evolution: A look into the past. Wiley–Blackwell, John Wiley & Sons Ltd., Publication, p. 317–334. Chichester, UK.

Jaramillo, C., Ochoa, D., Contreras, L., Pagani, M., Carvajal–Ortiz, H., Pratt, L.M., Krishnan, S., Cardona, A., Romero, M., Quiroz, L., Rodríguez, G., Rueda, M., De la Parra, F., Moron, S., Green, W., Bayona, G., Montes, C., Quintero, O., Ramírez, R., Mora, A., Schouten, S., Bermúdez, H., Navarrete, R.E., Parra, F., Alvarán, M., Osorno, J., Crowley, J.L., Valencia, V. & Vervoort, J. 2010b. Effects of rapid global warming at the Paleocene – Eocene boundary on Neotropical vegetation. Science, 330(6006): 957–961.

Jaramillo, C., Cadena, E. & Herrera, F. 2014a. Diversidad fósil en el valle de Cerrejón. In: Báez, L. & Trujillo, F. (editors), Biodiversidad en Cerrejón. Carbones de Cerrejón, Fundación Omacha, Fondo para la Acción Ambiental y la Niñez. p. 39–55. Bogotá.

Jaramillo, C., Moreno, E., Ramirez, V., da Silva, S., de la Barrera, A., de la Barrera, A., Sánchez, C., Morón, S., Herrera, F., Escobar, J., Koll, R., Manchester, S.R. & Hoyos, N. 2014b. Palynological record of the last 20 million years in Panama. In: Stevens, W.D., Montiel, O.M. & Raven, P.H. (editors), Paleobotany and biogeography: A Festschrift for Alan Graham in his 80th year. Missouri Botanical Garden Press, p. 134–251. Saint Louis, USA.

Jaramillo, C., Moreno, F., Hendy, F., Sánchez–Villagra, M. & Marty, D. 2015. Preface: La Guajira, Colombia: A new window into the Cenozoic Neotropical biodiversity and the Great American Biotic Interchange. Swiss Journal of Palaeontology, 134: 1–4.

Jaramillo, C., Montes, C., Cardona, A., Silvestro, D., Antonelli, A. & Bacon, C.D. 2017a. Comment (1) on "Formation of the Isthmus of Panama" by O'Dea et al. Science Advances, 3(6): 1–8.

Jaramillo, C., Romero, I., D'Apolito, C., Bayona, G., Duarte, E., Louwye, S., Escobar, J., Luque, J., Carrillo–Briceno, J., Zapata, V., Mora, A., Schouten, S., Zavada, M., Harrington, G., Ortiz, J. & Wesselingh, F. 2017b. Miocene flooding events of western Amazonia. Science Advances, 3(5): 1–11.

Jud, N.A., Nelson, C.W. & Herrera, F. 2016. Fruits and wood of Parinari from the early Miocene of Panama and the fossil record of Chrysobalanaceae. American Journal of Botany, 103(2): 277–289.

Kar, N., Garzione, C.N., Jaramillo, C., Shanahan, T., Carlotto, V., Pullen, A., Moreno, F., Anderson, V., Moreno, E. & Eiler, J. 2016. Rapid regional surface uplift of the northern Altiplano Plateau revealed by multiproxy paleoclimate reconstruction. Earth and Planetary Science Letters, 447: 33–47.

Kay, R.F., Madden, R.H., Cifelli, R.L. & Flynn, J.J., editors. 1997. Vertebrate paleontology in the Neotropics: The Miocene fauna of La Venta, Colombia. Smithsonian Institution Press, 608 p. Washington, D.C.

Keigwin, L.D. 1982. Isotopic paleoceanography of the Caribbean and East Pacific: Role of Panama uplift in late Neogene time. Science, 217(4557): 350–353.

Kemp, E.M. 1968. Probable angiosperm pollen from the British Barremian to Albian strata. Palaeontology, 11(3): 421–434.

Kennett, J.P. & Stott, L.D. 1991. Abrupt deep–sea warming, palaeoceanographic changes and benthic extinctions at the end of the Paleocene. Nature, 353: 225–229.

Krause, G.H., Winter, K., Krause, B., Jahns, P., García, M., Aranda, J. & Virgo, A. 2010. High–temperature tolerance of a tropical tree, Ficus insipida: Methodological reassessment and climate change considerations. Functional Plant Biology, 37(9): 890–900.

Kreft, H. & Jetz, W. 2007. Global patterns and determinants of vascular plant diversity. Proceedings of the National Academy of Sciences of the United States of America, 104(14): 5925–5930.

Lammertsma, E.I., Boer, H.J., Dekker, S.C., Dilcher, D.L., Lotter, A.F. & Wagner–Cremer, F. 2011. Global CO2 rise leads to reduced maximum stomatal conductance in Florida vegetation. Proceedings of the National Academy of Sciences of the United States of America, 108(10): 4035–4040.

Latrubesse, E.M., Cozzuol, M., da Silva–Caminha, S.A., Rigsby, C.A., Absy, M.L. & Jaramillo, C. 2010. The late Miocene paleogeography of the Amazon Basin and the evolution of the Amazon River system. Earth–Science Reviews, 99(3–4): 99–124.

Lefebvre, V., Donnadieu, Y., Sepulchre, P., Swingedou, D. & Zhang, Z. 2012. Deciphering the role of southern gateways and carbon dioxide on the onset of the Antarctic Circumpolar Current. Paleoceanography and Paleoclimatology, 27(4): 1–9.

Lehmann, C.E., Archibald, S.A., Hoffmann, W.A. & Bond, W.J. 2011. Deciphering the distribution of the savanna biome. New Phytologist, 191(1): 197–209.

Leigh, E.G., Davidar, P., Dick, C., Puyravaud, J., Terborgh, J., ter Steege, H. & Wright, S. 2004. Why do some tropical forests have so many species of trees? Biotropica, 36(4): 447–473.

Leigh, E.G., O'Dea, A. & Vermeij, G.J. 2013. Historical biogeography of the Isthmus of Panama. Biological Reviews, 89(1): 148–172.

Leighton, L.R. 2005. The latitudinal diversity gradient through deep time: Testing the ''age of the tropics'' hypothesis using Carboniferous productidine brachiopods. Evolutionary Ecology, 19(6): 563–581.

Leite, R.N., Kolokotronis, S.O., Almeida, F.C., Werneck, F., Rogers, D.S. & Weksler, M. 2014. In the wake of invasion: Tracing the historical biogeography of the South American cricetid radiation (Rodentia, Sigmodontinae). PLOS ONE, 9(6): 1–12.

Lemoigne, Y. 1984. Données nouvelles sur la paléoflore de Colombie. Geobios, 17(6): 667–690.

Lerdau, M.T. & Throop, H.L. 1999. Isoprene emission and photosynthesis in a tropical forest canopy: Implications for model development. Ecological Applications, 9(4): 1109–1117.[1109:IEAPIA]2.0.CO;2

Lewis, S.L., Malhi, Y. & Phillips, O.L. 2004. Fingerprinting the impacts of global change on tropical forests. Philosophical Transactions of the Royal Society of London Series B: Biological Sciences, 359(1443): 437–462.

Linares, O. 2004. Bioestratigrafía de la fauna de mamíferos de las formaciones Socorro, Urumaco y Codore (Mioceno medio–Plioceno temprano, de la región de Urumaco, Falcon, Venezuela. Paleobiología Neotropical, 1: 1–26.

Liu, Z., Pagani, M., Zinniker, D., DeConto, R., Huber, B.T., Brinkhuis, H., Shah, S.R., Leckie, R.M. & Pearson, A. 2009. Global cooling during the Eocene – Oligocene climate transition. Science, 323(5918): 1187–1190.

Lloyd, J. & Farquhar, G.D. 1994. 13C discrimination during CO2 assimilation by the terrestrial biosphere. Oecologia, 99(3–4): 201–215.

Lloyd, J. & Farquhar, G.D. 2008. Effects of rising temperatures and [CO2] on the physiology of tropical forest trees. Philosophical Transactions of the Royal Society B, 363(1498): 1811–1817.

Lupia, R., Lidgard, S. & Crane, P.R. 1999. Comparing palynological abundance and diversity: Implications for biotic replacement during the Cretaceous angiosperm radiation. Paleobiology, 25(3): 305–340.

Lüthi, D., Le Floch, M., Bereiter, B., Blunier, T., Barnola, J.M., Siegenthaler, U., Raynaud, D., Jouzel, J., Fischer, H., Kawamura, K. & Stocker, T.F. 2008. High–resolution carbon dioxide concentration record 650 000–800 000 years before present. Nature, 453: 379–382.

MacFadden, B.J. 2006a. Extinct mammalian biodiversity of the ancient New World tropics. Trends in Ecology & Evolution, 21(3): 157–165.

MacFadden, B.J. 2006b. North American Miocene land mammals from Panama. Journal of Vertebrate Paleontology, 26(3): 720–734.[720:NAMLMF]2.0.CO;2

MacFadden, B.J. 2009. Three–toed browsing horse Anchiterium (Echidae) from the Miocene of Panama. Journal of Paleontology, 83(3): 489–492.

MacFadden, B.J. & Higgins, P. 2004. Ancient ecology of 15–million–year–old browsing mammals within C3 plant communities from Panama. Oecologia, 140(1): 169–182.

MacFadden, B.J., Kirby, M.X., Rincon, A., Montes, C., Moron, S., Strong, N. & Jaramillo, C. 2010. Extinct peccary "Cynorca" Occidentale (Tayassuidae, Tayassuinae) from the Miocene of Panama and correlations to North America. Journal of Paleontology, 84(2): 288–298.

MacFadden, B.J., Foster, D.A., Rincón, A.F., Morgan, G.S. & Jaramillo, C. 2012. The New World tropics as a cradle of biodiversity during the early Miocene: Calibration of the centenario fauna from Panama. Geological Society of America Abstracts with Programs, 44, p. 163.

Magallón, S. & Castillo, A. 2009. Angiosperm diversification through time. American Journal of Botany, 96(1): 349–365.

Magallón, S., Crane, P.R. & Herendeen, P.S. 1999. Phylogenetic pattern, diversity, and diversification of eudicots. Annals of the Missouri Botanical Garden, 86(2): 297–372.

Martínez, C., Carvalho, M., Madriñan, S. & Jaramillo, C.A. 2015. A Late Cretaceous Piper (Piperaceae) from Colombia and diversification patterns for the genus. American Journal of Botany, 102(2): 273–289.

Martínez, J.I. 2009. La historia cenozoica del fenómeno de El Niño. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 33(129): 491–512.

McInerney, F.A. & Wing, S.L. 2011. The Paleocene – Eocene Thermal Maximum: A perturbation of carbon cycle, climate, and biosphere with implications for the future. Annual Review of Earth and Planetary Sciences, 39: 489–516.

Mejía–Velásquez, P.J. 2007. Floral composition of a Lower Cretaceous paleotropical ecosystem inferred from quantitative palynology. Master thesis, University of Florida, 85 p. Gainesville, USA.

Mejía–Velásquez, P., Dilcher, D., Jaramillo, C., Fortini, L. & Manchester, R.S. 2012. Palynological composition of a Lower Cretaceous South American tropical sequence: Climatic implications and diversity comparisons with other latitudes. American Journal of Botany, 99(11): 1819–1827.

Mikolajewicz, U., Maier–Reimer, E., Crowley, T.J. & Kim, K.Y. 1993. Effect of Drake and Panamanian gateways on the circulation of an ocean model. Paleoceanography and Paleoclimatology, 8(4): 409–426.

Miller, K.G., Kominz, M.A., Browning, J.V., Wright, J.D., Mountain, G.S., Katz, M.E., Sugarman, P.J., Cramer, B.S., Christie–Blick, N. & Pekar, S.F. 2005. The Phanerozoic record of global sea–level change. Science, 310(5752): 1293–1298.

Molnar, P. 2008. Closing of the Central American Seaway and the ice age: A critical review. Paleoceanography and Paleoclimatology, 23(2): 1–15.

Molnar, P. 2017. Comment (2) on "Formation of the Isthmus of Panama" by O'Dea et al. Science Advances, 3(6): 1–4.

Monnin, E., Indermühle, A., Dällenbach, A., Flückiger, J., Stauffer, B., Stocker, T.F., Raynaud, D. & Barnola, J.M. 2001. Atmospheric CO2 concentrations over the last glacial termination. Science, 291(5501): 112–114.

Montes, C., Bayona, G., Cardona, A., Buchs, D.M., Silva, C.A., Morón, S., Hoyos, N., Ramírez, D.A., Jaramillo, C. & Valencia, V. 2012a. Arc–continent collision and orocline formation: Closing of the Central American Seaway. Journal of Geophysical Research: Solid Earth, 117(B4): 25 p.

Montes, C., Cardona, A., McFadden, R.R., Moron, S., Silva, C.A., Restrepo–Moreno, S., Ramírez, D., Hoyos, N., Wilson, J., Farris, D.W., Bayona, G., Jaramillo, C., Valencia, V., Bryan, J. & Flores, J.A. 2012b. Evidence for middle Eocene and younger land emergence in Central Panama: Implications for isthmus closure. Geological Society of America Bulletin, 124(5–6): 780–799.

Montes, C., Cardona, A., Jaramillo, C., Pardo, A., Silva, J.C., Valencia, V., Ayala, C., Pérez–Ángel, L.C., Rodríguez–Parra, L.A., Ramírez, V. & Niño, H. 2015. Middle Miocene closure of the Central American Seaway. Science, 348(6231): 226–229.

Moore, M.J., Soltis, P.S., Bell, C.D., Burleigh, J.G. & Soltis, D.E. 2010. Phylogenetic analysis of 83 plastid genes further resolves the early diversification of eudicots. Proceedings of the National Academy of Sciences of the United States of America, 107(10): 4623–4628.

Moreno–Bernal, J.W., Head, J. & Jaramillo, C. 2016. Fossil crocodilians from the high Guajira Peninsula of Colombia: Neogene faunal change in northernmost South America. Journal of Vertebrate Paleontology, 36(3): 1–17.

Moreno, J.F., Hendy, A.J.W., Quiroz, L., Hoyos, N., Jones, D.S., Zapata, V., Zapata, S., Ballen, G.A., Cadena, E., Cárdenas, A.L., Carrillo–Briceño, J.D., Carrillo, J.D., Delgado–Sierra, D., Escobar, J., Martínez, J.I., Martínez, C., Montes, C., Moreno, J., Pérez, N., Sánchez, R., Suárez, C., Vallejo–Pareja, M.C. & Jaramillo, C. 2015. Revised stratigraphy of Neogene strata in the Cocinetas Basin, La Guajira, Colombia. Swiss Journal of Palaeontology, 134(1): 5–43.

Morgan, M.E., Kingston, J.D. & Marino, B.D. 1994. Carbon isotopic evidence for the emergence of C4 plants in the Neogene from Pakistan and Kenya. Nature, 367: 162–165.

Moritz, C., Patton, J.L., Schneider, C.J. & Smith, T.B. 2000. Diversification of rainforest faunas: An integrated molecular approach. Annual Review of Ecology and Systematics, 31: 533–563.

Muller–Landau, H.C. 2010. The tolerance–fecundity trade–off and the maintenance of diversity in seed size. Proceedings of the National Academy of Sciences of the United States of America, 107(9): 4242–4247.

Near, T.J., Dornburg, A., Kuhn, K.L., Eastman, J.T., Pennington, J.N., Patarnello, T., Zane, L., Fernández, D.A. & Jones, C.D. 2012. Ancient climate change, antifreeze, and the evolutionary diversification of Antarctic fishes. Proceedings of the National Academy of Sciences of the United States of America, 109(9): 3434–3439.

Nichols, D.J. & Johnson, K.G. 2008. Plants and the K–T boundary. Cambridge University Press, 292 p. Cambridge, UK.

Niu, S., Wu, M., Han, Y., Xia, J., Li, L. & Wan, S. 2008. Water–mediated responses of ecosystem carbon fluxes to climatic change in a temperate steppe. New Phytologist, 177(1): 209–219.

O'Dea, A., Lessios, H.A., Coates, A.H., Eytan, R., Restrepo–Moreno, S., Cione, A.L., Collins, L.S., De Queiroz, A., Farris, D.W., Norris, R.D., Stallard, R.F., Woodburne, M.O., Aguilera, O., Aubry, M., Berggren, W.A., Budd, A.F., Cozzuol, M.A., Coppard, S.E., Duque–Caro, H., Finnegan, S., Gasparini, G.M., Grossman, E.L., Johnson, K.G., Keigwin, L.D., Knowlton, N., Leigh, E.G., Leonard–Pingel, J.S., Marko, P.B., Pyenson, N.D., Rachello–Dolmen, P.G., Soibelzon, E., Soibelzon, L., Todd, J.A., Vermeij, G.J. & Jackson, J.B. 2016. Formation of the Isthmus of Panama. Science Advances, 2(8): 1–11.

Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D'Amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P. & Kassem, K.R. 2001. Terrestrial ecoregions of the world: A new map of life on Earth. BioScience, 51(11): 933–938.[0933:TEOTWA]2.0.CO;2

Ornelas, J.F., González, C., Espinosa de los Monteros, A., Rodríguez–Gómez, F. & García–Feria, L.M. 2013. In and out of Mesoamerica: Temporal divergence of Amazilia hummingbirds pre–dates the orthodox account of the completion of the Isthmus of Panama. Journal of Biogeography, 41(1): 168–181.

Pagani, M., Liu, Z.H., LaRiviere, J. & Ravelo, A.C. 2010. High Earth–system climate sensitivity determined from Pliocene carbon dioxide concentrations. Nature Geoscience, 3: 27–30.

Pardo–Trujillo, A. 2004. Paleocene – Eocene palynology and palynofacies from northeastern Colombia and western Venezuela. Doctorade thesis, Université de Liège, 103 p. Liège, Belgium.

Pardo–Trujillo, A. & Jaramillo, C. 2002. New palynostratigraphical data of NW South America Paleocene – Eocene of the Middle Magdalena Valley, Colombia. International Journal of Tropical Geology, Geography and Ecology, 26(1): 1–10.

Pardo–Trujillo, A., Jaramillo, C. & Oboh–Ikuenobe, F. 2003. Paleogene palynostratigraphy of the eastern Middle Magdalena Valley, Colombia. Palynology, 27(1): 155–178.

Peppe, D.J., Royer, D.L., Cariglino, B., Oliver, S.Y., Newman, S., Leight, E., Enikolopov, G., Fernández–Burgos, M., Herrera, F., Adams, J.M., Correa, E., Currano, E.D., Erickson, J.M., Hinojosa, L.F., Hoganson, J.W., Iglesias, A., Jaramillo, C.A., Johnson, K.R., Jordan, G.J., Kraft, N.J.B., Lovelock, E.C., Lusk, C.H, Niinemets, U., Peñuelas, J., Rapson, G., Wing, S.L. & Wright, I.J. 2011. Sensitivity of leaf size and shape to climate: Global patterns and paleoclimatic applications. New Phytologist, 190(3): 724–739.

Pérez–Consuegra, N., Cuervo–Gómez, A., Martínez, C., Montes, C., Herrera, F., Madriñán, S. & Jaramillo, C. 2017. Paleogene Salvinia (Salviniaceae) from Colombia and their paleobiogeographic implications. Review of Palaeobotany and Palynology, 246: 85–108.

Pérez–Consuegra, N., Parra, M., Jaramillo, C., Silvestro, D., Echeverri, S., Montes, C., Jaramillo, J.M. & Escobar, J. 2018. Provenance analysis of the Pliocene Ware Formation in the Guajira Peninsula, northern Colombia: Paleodrainage implications. Journal of South American Earth Sciences, 81: 66–77.

Pérez, M., Vallejo–Pareja, M.C., Carrillo, J.D. & Jaramillo, C. 2017. A new Pliocene capybara (Rodentia, Caviidae) from northern South America (Guajira, Colombia), and its implications for the Great American Biotic Interchange. Journal of Mammalian Evolution, 24(1): 111–125.

Pimiento, C., Griffin, J.N., Clements, C.F., Silvestro, D., Varela, S., Uhen, M. & Jaramillo, C. 2017. The Pliocene marine megafauna extinction and its impact on functional diversity. Nature Ecology & Evolution, 1: 1100–1106.

Pinto–Sánchez, N., Ibáñez, R., Madriñán, S., Sanjur, O., Bermingham, E. & Crawford, A.J. 2012. The Great American Biotic Interchange in frogs: Multiple and early colonization of Central America by the South American genus Pristimantis (Anura: Craugastoridae). Molecular Phylogenetics and Evolution, 62(3): 954–972.

Pons, D. 1988. Le Mesozoique de Colombie: Macroflores et microflores, Paris, Editions du Centre National de la Recherche Scientifique: Diffusion Presses du CNRS, Cahiers de paleontologie. Travaux de paliontologie est–africaine, 168 p.

Poveda, G., Waylen, P.R. & Pulwarty, R.S. 2006. Annual and inter–annual variability of the present climate in northern South America and southern Mesoamerica. Palaeogeography, Palaeoclimatology, Palaeoecology, 234(1): 3–27.

Quiroz, L.I. & Jaramillo, C. 2010. Stratigraphy and sedimentary environments of Miocene shallow to marginal marine deposits in the Urumaco Trough, Falcon Basin, western Venezuela. In: Sánchez–Villagra, M., Aguilera, O. & Carlini, A.A. (editors), Urumaco and Venezuelan paleontology: The fossil record of the northern Neotropics. Indiana University Press, p. 153–172. Bloomington, USA.

Ramírez, S.R., Gravendeel, B., Singer, R.B., Marshall, C.R. & Pierce, N.E. 2007. Dating the origin of the Orchidaceae from a fossil orchid with its pollinator. Nature, 448: 1042–1045.

Rangel, A., Moldowan, J.M., Nino, C., Parra, P. & Giraldo, B.N. 2002. Umir Formation: Organic geochemical and stratigraphic assessment as cosource for Middle Magdalena Basin oil, Colombia. American Association of Petroleum Geologists Bulletin, 86(12): 2069–2087.

Ravelo, A.C., Dekens, P.S. & McCarthy, M. 2006. Evidence for El Niño–like conditions during the Pliocene. Geological Society of America Today, 16(3): 4–11.<4:EFENLC>2.0.CO;2

Ricklefs, R.E. & Renner, S.S. 2012. Global correlations in tropical tree species richness and abundance reject neutrality. Science, 335(6067): 464–467.

Rincon, A., Bloch, J.I., Suárez, C., MacFadden, B.J. & Jaramillo, C. 2012. New floridatragulines (Mammalia, Camelidae) from the early Miocene Las Cascadas Formation, Panama. Journal of Vertebrate Paleontology, 32(2): 456–475.

Rincon, A., Bloch, J.I., MacFadden, B.J. & Jaramillo, C. 2013. First Central American record of Anthracotheriidae (Mammalia, Bothriodontinae) from the early Miocene of Panama. Journal of Vertebrate Paleontology, 33(2): 421–433.

Rincón–Martínez, D., Lamy, F., Contreras, S., Leduc, G., Bard, E., Saukel, C., Blanz, T., Mackensen, A. & Tiedemann, R. 2010. More humid interglacials in Ecuador during the past 500 kyr linked to latitudinal shifts of the equatorial front and the intertropical convergence zone in the eastern tropical Pacific. Paleoceanography and Paleoclimatology, 25(2): 1–15.

Rodríguez–Reyes, O., Falcon–Lang, H.J., Gasson, P., Collinson, M.E. & Jaramillo, C. 2014. Fossil woods (Malvaceae) from the lower Miocene (early to mid–Burdigalian) part of the Cucaracha Formation of Panama (Central America) and their biogeographic implications. Review of Palaeobotany and Palynology, 209: 11–34.

Rodríguez–Reyes, O., Gasson, P., Falcon–Lang, H.J. & Collinson, M.E. 2017a. Fossil legume woods of the Prioria–clade (subfamily Detarioideae) from the lower Miocene (early to mid–Burdigalian) part of the Cucaracha Formation of Panama (Central America) and their systematic and palaeoecological implications. Review of Palaeobotany and Palynology, 246: 44–61.

Rodríguez–Reyes, O., Gasson, P., Thornton, C.V., Falcon–Lang, H.J. & Jud, N.A. 2017b. Panascleroticoxylon crystallosa gen. et sp. nov.: A new Miocene malpiguialean tree from Panama. IAWA Journal, 38(4): 437–455.

Rosenzweig, M.L. 1995. Species diversity in space and time. Cambridge University Press, 460 p. Cambridge.

Roubik, D.W. & Camargo, J.M.F. 2011. The Panama microplate, island studies and relictual species of Melipona (Melikerria) (Hymenoptera: Apidae: Meliponini). Systematic Entomology, 37(1): 189–199.

Royer, D. 2006. CO2–forced climate thresholds during the Phanerozoic. Geochimica et Cosmochimica Acta, 70(23): 5665–5675.

Royer, D. 2010. Fossil soils constrain ancient climate sensitivity. Proceedings of the National Academy of Sciences of the United States of America, 107(2): 517–518.

Royer, D. 2016. Climate sensitivity in the geologic past. Annual Review of Earth and Planetary Sciences, 44: 277–293.

Royer, D., Pagani, M. & Beerling, D.J. 2011. Geologic constraints on Earth system sensitivity to CO2 during the Cretaceous and early Paleogene. Earth System Dynamic Discussions, 2: 211–240.

Royer, D. Pagani, M. & Beerling, D.J. 2012. Geobiological constraints on Earth system sensitivity to CO2 during the Cretaceous and Cenozoic. Geobiology, 10(4): 298–310.

Sacek, V. 2014. Drainage reversal of the Amazon River due to the coupling of surface and lithospheric processes. Earth and Planetary Science Letters, 401: 301–312.

Sage, R.F., Wedin, D.A. & Li, M. 1999. The biogeography of C4 photosynthesis: Patterns and controlling factors. In: Sage, R.F. & Monson, R.K. (editors), C4 plant biology. Academic Press, p. 313–373. San Diego, USA.

Sánchez–Villagra, M. 2006. Vertebrate fossils from the Neogene of Falcón state, Venezuela: Contributions on Neotropical palaeontology. Journal of Systematic Palaeontology, 4(3): 211.

Sánchez–Villagra, M. & Aguilera, O. 2006. Neogene vertebrates from Urumaco, Falcón state, Venezuela: Diversity and significance. Journal of Systematic Palaeontology, 4(3): 213–220.

Sánchez–Villagra, M., Aguilera, O. & Horovitz, I. 2003. The anatomy of the world's largest extinct rodent. Science, 301(5640): 1708–1710.

Sarmiento, G. 1992. Palinología de la Formación Guaduas–estratigrafía y sistemática. Boletín Geológico, 32(1–3): 45–126.

Scheyer, T.M., Aguilera, O.A., Delfino, M., Fortier, D.C., Carlini, A.A., Sánchez, R., Carrillo–Briceño, J.D., Quiroz, L. & Sánchez–Villagra, M.R. 2013. Crocodylian diversity peak and extinction in the late Cenozoic of the northern Neotropics. Nature Communications, 4(1907): 1–9.

Schuettpelz, E. & Pryer, K.M. 2009. Evidence for a Cenozoic radiation of ferns in an angiosperm–dominated canopy. Proceedings of the National Academy of Sciences of the United States of America, 106(27): 11200–11205.

Schulte, P., Alegret, L., Arenillas, I., Arz, J.A., Barton, P.J., Bown, P.R., Bralower, T.J., Christeson, G.L., Claeys, P., Cockell, C.S., Collins, G.S., Deutsch, A., Goldin, T.J., Goto, K., Grajales–Nishimura, J.M., Grieve, R.A.F., Gulick, S.P.S., Johnson, K.R., Kiessling, W., Koeberl, C., Kring, D.A., MacLeod, K.G., Matsui, T., Melosh, J., Montanari, A., Morgan, J.V., Neal, C.R., Nichols, D.J., Norris, R.D., Pierazzo, E., Ravizza, G., Rebolledo–Vieyra, M., Reimold, W.U., Robin, E., Salge, T., Speijer, R.P., Sweet, A.R., Urrutia–Fucugauchi, J., Vajda, V., Whalen, M.T. & Willumsen, P.S. 2010. The Chicxulub asteroid impact and mass extinction at the Cretaceous – Paleogene boundary. Science, 327(5970): 1214–1218.

Schultz, T.R. & Brady, S.G. 2008. Major evolutionary transitions in ant agriculture. Proceedings of the National Academy of Sciences of the United States of America, 105(14): 5435–5440.

Sepulchre, P., Sloan, L.C., Snyder, M. & Fiechter, J. 2009. Impacts of Andean uplift on the Humboldt Current system: A climate model sensitivity study. Paleoceanography and Paleoclimatology, 24(4): 1–11.

Sepulchre, P., Sloan, L.C. & Fluteau, F. 2010. Modelling the response of Amazonian climate to the uplift of the Andean mountain range. In: Hoorn, C. & Wesselingh, F.P. (editors), Amazonia: Landscape and species evolution: A look into the past. Wiley–Blackwell, John Wiley & Sons Ltd., Publication, p. 211–222. Chichester, UK.

Sepulchre, P., Arsouze, T., Donnadieu, Y., Dutay, J.C., Jaramillo, C., Le Bras, J., Martin, E., Montes, C. & Waite, A.J. 2014. Consequences of shoaling of the Central American Seaway determined from modeling Nd isotopes. Paleoceanography and Paleoclimatology, 29(3): 176–189.

Seton, M., Müller, R.D., Zahirovic, S., Gaina, C., Torsvik, T.H., Shephard, G., Talsma, A., Gurnis, M., Turner, M., Maus, S. & Chandler, M. 2012. Global continental and ocean basin reconstructions since 200 Ma. Earth–Science Reviews, 113(3–4): 212–270.

Shackleton, N.J., Backman, J., Zimmerman, H., Kent, D.V., Hall, M.A., Roberts, D.G., Schnitker, D., Baldauf, J.G., Desprairies, A., Homrighausen, R., Huddlestun, P., Keene, J.B., Kaltenback, A.J., Krumsiek, K.A.O., Morton, A.C., Murray, J.W. & Westberg–Smith, J. 1984. Oxygen isotope calibration of the onset of ice–rafting and history of glaciation in the North Atlantic region. Nature, 307: 620–623.

Shellito, C.J., Sloan, L.C. & Huber, M. 2003. Climate model sensitivity to atmospheric CO2 levels in the early – middle Paleogene. Palaeogeography, Palaeoclimatology, Palaeoecology, 193(1): 113–123.

Shephard, G.E., Müller, R.D., Liu, L. & Gurnis, M. 2010. Miocene drainage reversal of the Amazon River driven by plate–mantle interaction. Nature Geoscience, 3: 870–875.

Siegenthaler, U., Stocker, T.F., Monnin, E., Lüthi, D., Schwander, J., Stauffer, B., Raynaud, D., Barnola, J.M., Fischer, H., Masson–Delmotte, V. & Jouzel, J. 2005. Stable carbon cycle–climate relationship during the late Pleistocene. Science, 310(5752): 1313–1317.

Simpson, G.G. 1983. Splendid isolation: The curious history of South American mammals. Yale University Press, 275 p. New Haven, USA.

Slaughter, B.H. 1981. A new genus of geomyoid rodent from the Miocene of Texas and Panama. Journal of Vertebrate Paleontology, 1(1): 111–115.

Sloan, L.C. & Barron, E.J. 1992. A comparison of Eocene climate model results to quantified paleoclimatic interpretations. Palaeogeography, Palaeoclimatology, Palaeoecology, 93(3–4): 183–202.

Sloan, L.C. & Morrill, C. 1998. Orbital forcing and Eocene continental temperatures. Palaeogeography, Palaeoclimatology, Palaeoecology, 144(1–2): 21–35.

Sloan, L.C. & Rea, D.K. 1996. Atmospheric carbon dioxide and early Eocene climate: A general circulation modeling sensitive study. Palaeogeography, Palaeoclimatology, Palaeoecology, 119(3–4): 275–292.

Sloan, L.C. & Thomas, E. 1998. Global climate of the late Paleocene epoch: Modeling the circumstances associated with a climatic "event". In: Aubry, M.P., Lucas, S.G. & Berggren, W.A. (editors), Late Paleocene – early Eocene climatic and biotic events in the marine and terrestrial records. Columbia University Press, p. 138–157. New York.

Sloan, L.C., Walker, J.C. & Moore, T.C. 1995. Possible role of oceanic heat transport in early Eocene climate. Paleoceanography and Paleoclimatology, 10(2): 347–356.

Slot, M. & Winter, K. 2017. Photosynthetic acclimation to warming in tropical forest tree seedlings. Journal of Experimental Botany, 68(9): 2275–2284.

Smith, B.T., Amei, A. & Klicka, J. 2012. Evaluating the role of contracting and expanding rainforest in initiating cycles of speciation across the Isthmus of Panama. Proceedings of Royal Society of London Series B: Biological Sciences, 279(1742): 3520–3526.

Sole de Porta, N. 1971. Algunos géneros nuevos de polen procedentes de la Formación Guaduas (Maastrichtiense – Paleoceno) de Colombia. Studia Geologica, 2: 133–143.

Stoskopf, N. 1981. Understanding crop production: Upper Saddle River, Reston, Virginia. Reston Publishing Company, Inc. 433 p.

Stull, G.W., Herrera, F., Manchester, S., Jaramillo, C. & Tiffney, B.H. 2012. Fruits of an "Old World" tribe (Phytocreneae; Icacinaceae) from the Paleogene of North and South America. Systematic Botany, 37(3): 784–794.

Suárez, C., Forasiepi, A.M., Goin, F.J. & Jaramillo, C. 2016. Insights into the Neotropics prior to the Great American Biotic Interchange: New evidence of mammalian predators from the Miocene of northern Colombia. Journal of Vertebrate Paleontology, 36(1): p. 1–10.

Sucerquia, P. & Jaramillo, C. 2008. Lower Cretaceous floras from central Colombia. Palynology, 32: 271–272.

Sun, G. & Dilcher, D. 2002. Early angiosperms from the Lower Cretaceous of Jixi, eastern Heilongjiang, China. Review of Palaeobotany and Palynology, 121(2): 91–112.

Sun, G., Ji, Q., Dilcher, D.L., Zheng, S., Nixon, K.C. & Wang, X. 2002. Archaefructaceae, a new basal angiosperm family. Science, 296(5569): 899–904.

Sun, G., Dilcher, D., Wang, H. & Chen, Z. 2011. A eudicot from the Early Cretaceous of China. Nature, 471: 625–628.

Tewksbury, J.J., Huey, R.B. & Deutsch, C.A. 2008. Putting the heat on tropical animals. Science, 320(5881): 1296–1297.

Thomas, E. & Shackleton, N.J. 1996. The Paleocene – Eocene benthic foraminiferal extinction and stable isotope anomalies. In: Knox, R., Corfield, R.M. & Dunay, R.E. (editors), Correlations of the early Paleogene in Northwest Europe: An overview. Geological Society of London, Special Publication 101, p. 401–441.

Thomas, W.W. 1999. Conservation and monographic research on the flora of tropical America. Biodiversity & Conservation, 8(8): 1007–1015.

Toivonen, T., Maki, S. & Kalliola, R. 2007. The riverscape of western Amazonia—A quantitative approach to the fluvial biogeography of the region. Journal of Biogeography, 34(8): 1374–1387.

Tripati, A., Roberts, C. & Eagle, R. 2009. Coupling of CO2 and ice sheet stability over major climate transitions of the last 20 million years. Science, 326(5958): 1394–1397.

Ufnar, D.F., González, L.A., Ludvigson, G.A., Brenner, R.L. & Witzke, B.J. 2002. The mid–Cretaceous water bearer: Isotope mass balance quantification of the Albian hydrologic cycle. Palaeogeography, Palaeoclimatology, Palaeoecology, 188(1–2): 51–71.

Ufnar, D.F., González, L.A., Ludvigson, G.A., Brenner, R.L. & Witzke, B.J. 2004. Evidence for increased latent heat transport during the Cretaceous (Albian) greenhouse warming. Geology, 32(12): 1049–1052.

Ufnar, D.F., Ludvigson, G.A., González, L.A. & Gröcke, D.R. 2008. Precipitation rates and atmospheric heat transport during the Cenomanian greenhouse warming in North America: Estimates from a stable isotope mass–balance model. Palaeogeography, Palaeoclimatology, Palaeoecology, 266(1–2): 28–38.

Uno, K.T., Cerling, T.E., Harris, J.M., Kunimatsu, Y., Leakey, M.G., Nakatsukaka, M. & Nakaya, H. 2011. Late Miocene to Pliocene carbon isotope record of differential diet change among east African herbivores. Proceedings of the National Academy of Sciences of the United States of America, 108(16): 6509–6514.

van der Hammen, T. 1958. Estratigrafía del Terciario y Maastrichtiano continentales y tectogénesis de los Andes colombianos. Boletín Geológico, 6(1–3): 67–128.

van der Hammen, T. 1989. History of the montane forests of the northern Andes. Plant Systematics and Evolution, 162(1–4): 109–114.

van der Hammen, T. 1995. Plioceno y Cuaternario del altiplano de Bogotá y alrededores. Análisis Geográficos, 24: 1–142.

van der Hammen, T. 2003. Neógeno y Cuaternario del altiplano de Bogotá y alrededores. Análisis Geográficos, 26: 101–120.

van der Hammen, T. & Hooghiemstra, H. 2000. Neogene and Quaternary history of vegetation, climate, and plant diversity in Amazonia. Quaternary Science Reviews, 19(8): 725–742.

van der Hammen, T., Werner, J.H. & van Dommelen, H. 1973. Palynological record of the upheaval of the northern Andes: A study of the Pliocene and lower Quaternary of the Colombian Eastern Cordillera and the early evolution of its high–Andean biota. Review of Palaeobotany and Palynology, 16(1–2): 1–122.

Wallis, G.P., Waters, J.M., Upton, P. & Craw, D. 2016. Transverse alpine speciation driven by glaciation. Trends in Ecology & Evolution, 31(12): 916–926.

Wang, H., Moore, M.J., Soltis, P.S., Bell, C.D., Brockington, S.F., Alexandre, R., Davis, C.C., Latvis, M., Manchester, S. & Soltis, D.E. 2009. Rosid radiation and the rapid rise of angiosperm–dominated forests. Proceedings of the National Academy of Sciences of the United States of America, 106(10): 3853–3858.

van Waveren, I.M., van Konijnenburg–van Cittert, J.H.A., van der Burgh, J. & Dilcher, D.L. 2002. Macrofloral remains from the Lower Cretaceous of the Leiva region (Colombia). Scripta Geologica, 123: 1–39.

Webb, S.D. 1976. Mammalian faunal dynamics of the Great American Interchange. Paleobiology, 2(3): 220–234.

Webb, S.D. 1978. A history of savanna vertebrates in the New World. Part II: South America and the great interchange. Annual Review of Ecology and Systematics, 9: 393–426.

Webb, S.D. 1994. Successful in spite of themselves. Natural History, 4: 50–53.

Webb, S.D. 1995. Biological implications of the middle Miocene Amazon seaway. Science, 269(5222): 361–362.

Webb, S.D. 2006. The Great American Biotic Interchange: Patterns and processes. Annals of the Missouri Botanical Garden, 93(2): 245–257.[245:TGABIP]2.0.CO;2

Westerhold, T., Röhl, U., McCarren, H.K. & Zachos, J.C. 2009. Latest on the absolute age of the Paleocene – Eocene Thermal Maximum (PETM): New insights from exact stratigraphic position of key ash layers +19 and –17. Earth and Planetary Science Letters, 287(3–4): 412–419.

Whitmore, F.C. & Stewart, R.H. 1965. Miocene mammals and Central American seaways: Fauna of the Canal zone indicates separation of Central and South America during most of the Tertiary. Science, 148(3667): 180–185.

Wijmstra, T.A. & van der Hammen, T. 1966. Palynological data on the history of tropical savannas in northern South America. Leidse Geologische Mededelingen, 38(1): 71–90.

Wijninga, V.M. 1996. Paleobotany and palynology of Neogene sediments from the High Plain of Bogota (Colombia). Evolution of the Andean flora from a paleoecological perspective. Doctorade thesis, University of Amsterdam, 370 p. Amsterdam, the Netherlands.

Wikström, N., Savolainen, V. & Chase, M.W. 2001. Evolution of the angiosperms: Calibrating the family tree. Proceedings of the Royal Society London B: Biological Sciences, 268(1482): 2211–2220.

Williams, J.H. 2008. Novelties of the flowering plant pollen tube underlie diversification of a key life history stage. Proceedings of the National Academy of Sciences of the United States of America, 105(32): 11259–11263.

Wing, S.L. & Boucher, L. 1998. Ecological aspects of the Cretaceous flowering plant radiation. Annual Review of Earth and Planetary Sciences, 26: 379–421.

Wing, S.L., Hickey, L.J. & Swisher, C.C. 1993. Implications of an exceptional fossil flora for Late Cretaceous vegetation. Nature, 363: 342–344.

Wing, S.L., Harrington, G.J., Smith, F., Bloch, J.I., Boyer, D.M. & Freeman, K.H. 2005. Transient floral change and rapid global warming at the Paleocene – Eocene boundary. Science, 310(5750): 993–996.

Wing, S.L., Herrera, F., Jaramillo, C., Gómez–Navarro, C., Wilf, P. & Labandeira, C.C. 2009. Late Paleocene fossils from the Cerrejón Formation, Colombia, are the earliest record of Neotropical rainforest. Proceedings of the National Academy of Sciences of the United States of America, 106(44): 18627–18632.

Woodburne, M.O. 2010. The Great American Biotic Interchange: Dispersals, tectonics, climate, sea level and holding pens. Journal of Mammalian Evolution, 17(4): 245–264.

Wright, S., Keeling, J. & Gillman, L. 2006. The road from Santa Rosalia: A faster tempo of evolution in tropical climates. Proceedings of the National Academy of Sciences of the United States of America, 103(20): 7718–7722.

Wright, J.S., Fu, R., Worden, J.R., Chakraborty, S., Clinton, N.E., Risi, C., Sun, Y. & Yin, L. 2017. Rainforest–initiated wet season onset over the southern Amazon. Proceedings of the National Academy of Sciences of the United States of America, 114(32): 8481–8486.

Zachos, J.C., Pagani, M., Sloan, L., Thomas, E. & Billups, K. 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science, 292(5517): 686–693.

Zachos, J.C., Wara, M.W., Bohaty, S., Delaney, M.L., Petrizzo, M.R., Brill, A., Bralower, T.J. & Premoli–Silva, I. 2003. A transient rise in tropical sea surface temperature during the Paleocene – Eocene Thermal Maximum. Science, 302(5650): 1551–1554.