An evening bat (Chiroptera: Vespertilionidae) from the late Early Eocene of France, with comments on the antiquity of modern bats

Suzanne J. Hand, Bernard Sigé, Michael Archer and Karen H. Black

Cite this article: Hand S. J., Sigé B., Archer M., Black K. H., 2016. An evening bat (Chiroptera: Vespertilionidae) from the late Early Eocene of France, with comments on the antiquity of modern bats. Palaeovertebrata 40 (2)-e2. doi: 10.18563/pv.40.2.e2

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Abstract

Bats are among the most numerous and widespread mammals today, but their fossil record is comparatively meagre and their early evolution poorly understood. Here we describe a new fossil bat from dental remains recovered from late Early Eocene sediments at Prémontré, northern France. This 50 million-year-old bat exhibits a mosaic of plesiomorphic and apomorphic dental features, including the presence of three lower premolars, a single-rooted p3, short p4 with metaconid, myotodont lower molars and a tall coronoid process of the dentary. This combination of features suggests it is an early member of Vespertilionidae, today’s most speciose and geographically widespread bat family. The Prémontré bat has bearing on hypotheses about the origins of vesper or evening bats (Family Vespertilionidae), as well as crown-group chiropterans.


  

Bibliography

Adnet, S., Cappetta, H., 2008. New fossil triakid sharks from the early Eocene of Prémontré, France, and comments on fossil record of the family. Acta Palaeontologica Polonica 53, 433-448. http://dx.doi.org/10.4202/app.2008.0306
 
Aubry, M.-P., Thiry, M., Dupuis, C., Berggren, W., 2005. The Sparnacian deposits of the Paris Basin: a lithostratigraphic classification. Stratigraphy 2, 65-100.
 
Bininda-Emonds, O. R. P., Cardillo, M., Jones, K. E., MacPhee, R. D. E., Beck, R. M. D., Grenyer, R., Price, S. A., Vos, R. A., Gittleman, J. L., Purvis, A., 2007. The delayed rise of present-day mammals. Nature 446, 507-512. http://dx.doi.org/10.1038/nature05634
 
Cappetta, H., 1992. Carcharhiniformes nouveaux (Chondrichthyes, Neoselachii) de l’Yprésien du Bassin de Paris. Geobios 25, 639-646.

Collinson, M. E., 2000. Cenozoic evolution of modern plant communities and vegetation. In: Culver. S. J., Rawson, P. F. (Eds.), Biotic Response to Global Change. Cambridge University Press, Cambridge, pp. 223-243.
 
Czaplewski, N. J., G. S. Morgan., 2001. A new vespertilionid bat (Mammalia: Chiroptera) from the Early Miocene (Hemingfordian) of Florida. Journal of Vertebrate Paleontology 20, 736-742.
 
Czaplewski, N. J., Morgan, G. S., 2012. New basal noctilionoid bats (Mammalia: Chiroptera) from the Oligocene of subtropical North America. In: Gunnell, G  F., Simmons, N. B. (Eds.), Evolutionary History of Bats. Cambridge University Press, Boston, pp. 162-209.
 
Dégrémont, E., Duchaussois, F., Hautefeuille, F., Laurain, P., Louis, P., Tetu, R., 1985. Paléontologie: découverte d’un gisement du Cuisien tardif à Prémontré (Aisne). Bulletin d’Information des Géologues du Bassin de Paris 22, 11-18.
 
Eick, G. N., Jacobs, D. S., Matthee, C. A., 2005. A nuclear DNA phylogenetic perspective on the evolution of echolocation and historical biogeography of extant bats (Chiroptera). Molecular Biology and Evolution 22, 1869-1886.
 
Eiting, T. P., Gunnell, G. F., 2009. Global completeness of the bat fossil record. Journal of Mammalian Evolution 16, 151-173. http://dx.doi.org/ 10.1007/s10914-009-9118-x
 
Escarguel, G., 1999. Les rongeurs de l’Eocène inférieur et moyen d’Europe occidentale: systématique, phylogénie, biochronologie et paléobiogéographie des niveaux-repères MP 7 à MP 14. Palaeovertebrata 28, 89-351.
 
Giannini, N. P., Gunnell, G. F., Habersetzer, J., Simmons, N. B., 2012. Early evolution of body size. In: Gunnell, G. F., Simmons, N. B., (Eds.), Evolutionary History of Bats Fossils, Molecules and Morphology. Cambridge University Press, Boston, pp. 530-555.
 
Godinot, M., 2014. Fossil record of the Primates from the Paleocene to the Oligocene. In: Henke, W, Tattersal, I. (Ed.), Handbook of Paleoanthropology. Springer-Verlag, Berlin, pp. 1-102.
 
Godinot, M., Russell, D. E., Louis, P., 1992. Oldest known Nannopithex (Primates, Omomyiformes) from the early Eocene of France. Folia Primatologia 58, 32-40.
 
Gunnell, G. F., Simmons, N. B., 2005. Fossil evidence and the origin of bats. Journal of Mammalian Evolution 12, 209-246.
 
Gunnell, G. F., Simons, E. L., Seiffert, E. R., 2008. New bats (Mammalia: Chiroptera) from the late Eocene and early Oligocene, Fayum Depression, Egypt. Journal of Vertebrate Paleontology 28, 1-11.
 
Gunnell, G. F., Eiting, T. P., Simons, E. L, 2012. African Vespertilionoidea (Chiroptera) and the antiquity of Myotinae. In: Gunnell, G. F., Simmons, N. B. (Eds.) Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Boston, pp. 252-266.
 
Gunnell, G. F., Worsham, S. R., Seiffert, E. R., Simons, E. L., 2009. Vampyravus orientalis Schlosser (Chiroptera) from the early Oligocene of Egypt – body mass, humeral morphology and affinities. Acta Chiropterologica 11, 271-278.
 
Gunnell, G. F., Simmons, N. B., Seiffert, E. R., 2014. New Myzopodidae (Chiroptera) from the Late Paleogene of Egypt: emended family diagnosis and biogeographic origins of Noctilionoidea. PLoS ONE 9(2): e86712. http://dx.doi.org/10.1371/journal.pone.0086712
 
Gunnell, G. F., Habersetzer, J., Schlosser-Sturm, E., Simmons, N. B., Smith, T., 2011. Primitive chiropteran teeth: the complete dentition of the Messel bat Archaeonycteris trigonodon. In: Lehmann, T., Schaal, S. F. K. (Eds.), The World at the Time of Messel. HC Volker Mosbrugger, Senckenberg Research Institute and Natural History Museum, Frankfurt, pp. 73-76.
 
Habersetzer, J., Storch, G., 1987. Klassifikation und funktionelle Flügelmorphologie paläogener Fledermäuse (Mammalia, Chiroptera). Courier Forschungsinstitut Senckenberg 91, 117-150.
 
Habersetzer, J., Richter, G., Storch, G., 1994. Paleoecology of early middle Eocene bats from Messel, FRG. Aspects of flight, feeding and echolocation. Historical Biology 8, 235-260.
 
Hand, S.J., 1990.  First Tertiary molossid (Microchiroptera: Molossidae) from Australia: its phylogenetic and biogeographic implications. Memoirs of the Queensland Museum 28, 175-192.
 
Hand, S.J., Sigé, B., Maitre, E., 2012. Necromantis Weithofer, 1887, large carnivorous Middle and Late Eocene bats from the French Quercy Phosphorites: new data and unresolved relationships. In: Gunnell, G. F., Simmons, N. B. (Eds.) Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Boston, pp. 210-251.
 
Hand, S. J., Novacek, M., Godthelp, H., Archer, M., 1994. First Eocene bat from Australia. Journal of Vertebrate Paleontology 14, 375-381.
 
Hand, S.J., Murray, P.F., Megirian, D., Archer, M., Godthelp, H., 1998. Mystacinid bats (Microchiroptera) from the Australian Tertiary. Journal of Paleontology 72, 538-545.
 
Hand, S. J., Sigé, B., Archer, M., Gunnell, G. F., Simmons, N. B., 2015. A new early Eocene (Ypresian) bat from Pourcy, Paris Basin, France, with comments on patterns of diversity in the earliest chiropterans. Journal of Mammalian Evolution 22, 343-354. http://dx.doi.org/10.1007/s10914-015-9286-9
 
Harrison D. L., Hooker, J. J., 2010. Late middle Eocene bats from the Creechbarrow Limestone Formation, Dorset, south England with description of a new species of Archaeonycteris (Chiroptera: Archaeonycteridae). Acta Chiropterologica 12, 1-18.
 
Hoofer, S. R., Van Den Bussche, R. A., 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5, 1-63.
 
Hooker, J. J., 1996. A primitive emballonurid bat (Chiroptera, Mammalia) from the earliest Eocene of England. Palaeovertebrata 25, 287-300.
 
Horáček, I., 2001. On the early history of vespertilionid bats in Europe: the Lower Miocene record from the Bohemian Massif. Lynx 32, 123-154.
 
Horáček, I., Fejfar, O., Hulva, P., 2006. A new genus of vespertilionid bat from the Early Miocene of Jebel Zelten, Libya, with comments on Scotophilus and early history of vespertilionid bats (Chiroptera). Lynx 37, 131-150.
 
Huyghe, D., Lartaud, F., Emmanuel, L., Merle, D., Renard, M., 2015. Palaeogene climate evolution in the Paris Basin from oxygen stable isotope (d18O) compositions of marine molluscs. Journal of the Geological Society 172, 576-587. http://dx.doi.org/10.1144/jgs2015-016
 
Jones, K. E., Bininda-Emonds, O. R. P., Gittleman, J. L., 2005. Bats, clocks, and rocks: diversification patterns in Chiroptera. Evolution 59, 2243-2255.
 
Lack, J. B., Roehrs, Z. P., Stanley, C. E., Ruedi, M., Van Den Bussche, R. A., 2010. Molecular phylogenetics of Myotis indicate familial-level divergence for the genus Cistugo (Chiroptera). Journal of Mammalogy 91, 976-992. http://dx.doi.org/10.1644/09-MAMM-A-192.1
 
Lecomte, G., 1994. Étude paléontologique et sédimentologique de l’Yprésien de l’Est du Bassin de Paris. PhD, Université de Paris VI, Paris.
 
Legendre, S., 1984.  Étude odontologique des représentants actuels du groupe Tadarida (Chiroptera, Molossidae). Implications phylogéniques, systématiques et zoogéographiques.  Revue suisse de Zoologie 91, 399-442.
 
Legendre, S., 1985. Molossidés (Mammalia, Chiroptera) cénozoïques de l’Ancien et du Nouveau Monde : statut systématique; intégration phylogénique des données. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 170, 205-27.
 
Louis, P., 1996. Recherches de mammifères paléogènes dans les départements de l’Ainse et de la Marne pendant la deuxième moitié du vingtième siècle. Palaeovertebrata 25, 83-113.
 
Maitre, E., 2014. Western European middle Eocene to lower Oligocene Chiroptera – systematics, phylogeny and palaeoecology based on new material from the Quercy (France). Swiss Journal of Palaeontology 133, 141-242.
 
Marandat, B., 1991. Mammifères de l’Ilerdien moyen (Eocène inférieur) des Corbières et du Minervois (Bas-Languedoc, France). Systématique, biostratigraphie, corrélations. Palaeovertebrata 20, 55-144.
 
Menu, H., 1985. Morphotypes dentaires actuels et fossiles des chiroptères Vespertilioninés. 1ère partie: étude des morphologies dentaires. Palaeovertebrata 15, 71-128.
 
Menu, H., 1987. Morphotypes dentaires actuels et fossiles des chiroptères Vespertilioninés, 2ème partie: implications systématiques et phylogéniques. Palaeovertebrata 17, 77-150.
 
Menu, H., Sigé, B., 1971. Nyctalodontie et myotodontie, importants caractères de grades évolutifs chez les chiroptères entomophages. Comptes Rendus de l’Académie des Sciences Paris 272, 1735-1738.
 
Menzel, M. A., Carter, T. C., Chapman, B. R., Laerm, J., 1998. Quantitative comparison of tree roosts used by red bats (Lasiurus borealis) and Seminole bats (L. seminolus). Canadian Journal of Zoology 76, 630-634.
 
Meredith, R. W., Jane ka, J. E., Gatesy, J., Ryder, O. A., Fisher, C. A., Teeling, E. C., Goodbla,  A., Eizirik, E., Simão, T. L. L., Stadler, T., Rabosky, D. L., Honeycutt,  R. L., Flynn, J. J., Ingram, C. M., Steiner, C., Williams, T. L., Robinson, T. J., Burk-Herrick, A., Westerman, M., Ayoub, N. A., Springer, M. S., Murphy, W. J., 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334, 521-524. http://dx.doi.org/10.1126/science.1211028
 
Miller-Butterworth, C. M., Murphy, W. J., O’Brien, S. J., Jacobs, D. S., Springer, M. S., Teeling, E. C., 2007. A family matter: conclusive resolution of the taxonomic position of the long-fingered bats, Miniopterus. Molecular Biology and Evolution 24, 1553-1561.
 
Morgan, G. S., Czaplewski, N. J., 2003. A new bat (Chiroptera: Natalidae) from the Early Miocene of Florida, with comments on natalid phylogeny. Journal of Mammalogy 84, 729-752.
 
Morgan, G. S., Czaplewski, N. J., 2012. Evolutionary history of the Neotropical Chiroptera: the fossil record. In: Gunnell, G. F., Simmons, N. B., (Eds.), Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Boston, pp. 105-161.
 
Norberg, U. M., 1989. Ecological determinates of bat wing shape and echolocation call structure with implications for some fossil bats. In:  Hanák, V., Horácek, I., Gaisler, J. (Eds.), European Bat Research 1987. Charles University Press, Prague, pp. 197-211.
 
Norberg, U. M., Rayner, J. M., 1987. Ecological morphology and flight in bats (Mammalia; Chiroptera): wing adaptations, flight performance, foraging strategy and echolocation. Philosophical Transactions of the Royal Society B 316, 335-427.
 
O’Leary, M. A., Bloch, J. I., Flynn, J. J., Gaudin, T. J., Giallombardo, A., Giannini, N. P., Goldberg, S. L., Kraatz, B. P., Luo, Z.-X., Meng, J., Ni, X., Novacek, M. J., Perini, F.A., Randall, Z. S., Rougier, G. W., Sargis, E. J., Silcox, M. T., Simmons, N. B., Spaulding, M., Velazco, P. M., Weksler, M., Wible, J. R., Cirranello, A. L., 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science 339, 662-667. http://dx.doi.org/10.1126/science.1229237
 
Ravel, A., Marivaux, L., Tabuce, R., Adaci, M., Mahboubi, M., Mebrouk, F., Bensalah, M., 2011. The oldest African bat from the early Eocene of El Kohol (Algeria). Naturwissenshaften 98, 397–405. http://dx.doi.org/10.1007/s00114-011-0785-0
 
Ravel, A., Marivaux, L., Tabuce, R., Haj Ali, M. B.,  Essid, E. M., Vianey-Liaud, M.,  2012. A new large philisid (Mammalia, Chiroptera, Vespertilionoidea) from the late Early Eocene of Chambi, Tunisia. Palaeontology 55, 1035-1041. http://dx.doi.org/10.1111/j.1475-4983.2012.01160.x
 
Ravel, A., Marivaux, L., Qi, T., Wang, Y. O. Beard, K. C., 2014. New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province, Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia. Zoologica Scripta 43, 1-23. http://dx.doi.org/10.1111/zsc.12027
 
Ravel, A., Adaci, M., Bensalah, M., Mahboubi, M., Mebrouk, F., Essid, E. M., Marzougui, W., Khayati Ammar, H., Charruault, A.-L., Lebrun, R., Tabuce, R., Vianey-Liaud, M., Marivaux, L., 2015. New philisids (Mammalia, Chiroptera) from the early–middle Eocene of Algeria and Tunisia: new insight into the phylogeny, palaeobiogeography and palaeoecology of the Philisidae. Journal of Systematic Palaeontology 13, 691-709. http://dx.doi.org/10.1080/14772019.2014.941422
 
Revilliod, P., 1920. Contribution à l’étude des chiroptères des terrains tertiaires. 2.  Mémoires de la Société paléontologique Suisse 44, 63-129.

Roehrs, Z. P., Lack, J. B., Van Den Bussche, R. A., 2010. Tribal phylogenetic relationships within Vespertilioninae (Chiroptera: Vespertilionidae) based on mitochondrial and nuclear sequence data. Journal of Mammalogy 91, 1073-1092. http://dx.doi.org/10.1644/09-MAMM-A-325.1
 
Rose, K. D., Holbrook, L. T., Rana, R. S., Kumar, K., Jones, K. E., Ahrens, H. E., Missiaen, P., Sahni, A., Smith, T., 2014. Early Eocene fossils suggest that the mammalian order Perissodactyla originated in India.  Nature Communications 5: 5570. http://dx.doi.org/10.1038/ncomms6570
 
Russell, D. E., Sigé, B., 1970. Révision des chiroptères lutétiens de Messel (Hesse, Allemagne). Palaeovertebrata 3, 83-182.
 
Russell, D. E., Galoyer, A., Louis, P., Gingerich, P. D., 1988. Nouveaux vertébrés sparnaciens du Conglomérat de Meudon à Meudon, France. Comptes Rendus de l’Académie des Sciences Paris séries II 307, 429-433.
 
Scotese, C. R., 2006. PALEOMAP Project (http://www.scotese.com).
 
Sigé, B., 1974. Données nouvelles sur le genre Stehlinia (Vespertilionoidea, Chiroptera) du Paléogène d’Europe. Palaeovertebrata 6, 253-272.
 
Sigé, B., 1985. Les chiroptères Oligocènes du Fayum, Egypte. Geologica et Palaeontologica, 19, 161-89.
 
Sigé, B., 1991 Rhinolophoidea et Vespertilionoidea (Chiroptera) du Chambi (Eocène inférieur de Tunisie). Aspects biostratigraphique, biogéographique et paléoécologique de l’origine des chiroptères modernes. Neues Jahrbuch für Geologie und Paläontologie 182, 355-376.
 
Sigé, B., 1995. Le Garouillas et les sites contemporains (Oligocène, MP 25) des phosphorites du Quercy, Lot, Tarn & Garonne, France, et leurs faunes des vertébrés. 5 : Chiroptères. Palaeontographica A 236, 77-124.
 
Sigé, B., 1997. Les remplissages karstiques polyphasés (Eocène, Oligocène, Pliocène) de Saint-Maximin (phosphorite du Gard) et leur apport à la connaissance des faunes européennes, notamment pour l’Eocène moyen (MP 13). 3-Systématique: euthériens entomophages. In: Aguilar, J.-P., Legendre, R.-S., Michaux, J. (Eds.), Actes du Congres BiochroM’97. Mémoires et Travaux de l’Ecole pratique des Hautes Etudes, Institut de Montpellier, Montpellier, pp. 737-750.
 
Sigé, B., Maitre, E., Hand, S. J., 2012. Necromantodonty: the primitive condition of molar morphology in bats.  In: Gunnell, G. F., Simmons, N. B. (Eds.) Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Boston, pp. 456–469.
 
Simmons, N. B., 2005a. Order Chiroptera. In: Wilson, D. E., Reeder, D. M. (Eds.) Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd ed. Smithsonian Institution Press, Washington DC, pp 312-529.
 
Simmons, N. B., 2005b. An Eocene big bang for bats. Science 307, 527-528. 
 
Simmons, N. B., Geisler, J. H., 1998. Phylogenetic relationships of Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx to extant bat lineages, with comments on the evolution of echolocation and foraging strategies in Microchiroptera. Bulletin of the American Museum of Natural History 235, 1-182.
 
Simmons, N. B., Seymour, K. L., Habersetzer, J., Gunnell, G. F., 2008. Primitive early Eocene bat from Wyoming and the evolution of flight and echolocation. Nature 451, 818-822.
 
Slaughter, B. H., 1970. Evolutionary trends of chiropteran dentitions. In:, Slaughter, B. H., Walton, D. W. (Eds.), About Bats. Southern Methodist University Press, Dallas, pp. 51-83.
 
Smith, F. A., Lyons, S. K., Morgan, E. K., Jones, K. E., Kaufman, D. M., Dayan, T., Marquet, P. A., Brown, J. H., Haskell, J. P., 2004. Body mass of Late Quaternary mammals. Ecological Archives E084-094. Ecology 84, 3403.
 
Smith, T., Rana, R. S., Missiaen, P., Rose, K.D., Sahni, A., Singh, H., Singh, L., 2007. High bat (Chiroptera) diversity in the early Eocene of India. Naturwissenschaften 94, 1003-1009.
 
Smith, T., Habersetzer, J., Simmons, N. B., Gunnell, G. F., 2012. Systematics and paleobiogeography of early bats. In: Gunnell, G. F., Simmons, N. B. (Eds.) Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Boston, pp 23-66.
 
Smith, T., Solé, F., Missiaen, P., Rana, R. S., Kumar, K., Sahni, A., Rose, K. D., 2015. First Early Eocene tapiroid from India and its implication for the paleobiogeographic origin of perissodactyls.  Palaeovertebrata 39 (2), e5. http://dx.doi.org/10.18563/pv.39.2.e5
 
Stadelmann, B., Lin, L.-K., Kunz, T. H., Ruedi, M., 2007. Molecular phylogeny of New World Myotis (Chiroptera, Vespertilionidae) inferred from mitochondrial and nuclear DNA genes. Molecular Phylogenetics and Evolution 43, 32-48. http://dx.doi.org/10.1016/j.ympev.2006.06.019
 
Steurbaut, E. 1998. High-resolution holostratigraphy of middle Paleocene to early Eocene strata in Belgium and adjacent areas. Palaeontographica A 247, 91-156.
 
Storch, G., Sigé, B., Habersetzer, J., 2002. Tachypteron franzeni n. gen., n. sp., earliest emballonurid bat from the Middle Eocene of Messel (Mammalia, Chiroptera). Paläontologische Zeitschrift 76, 189-99.
 
Storer, J. E., 1984. Mammals of the Swift Current Creek local fauna (Eocene: Uintan), Saskatchewan. Natural History Contributions (Saskatchewan Culture and Recreation) 7, 1-158. 
 
Sudre, J., Erfurt, J. 1996. Les artiodactyles du gisement Yprésien terminal de Prémontré (Aisne, France) In: Godinot, M., Gingerich, P. D.  (Eds), Volume Jubilaire D.E. Russell. Palaeovertebrata 25, 391-414.
 
Tabuce, R., Antunes, M. T., Sigé, B., 2009. A new primitive bat from the earliest Eocene of Europe. Journal of Vertebrate Paleontology 29, 627-630.
 
Teeling, E. C., Springer, M. S., Madsen, O., Bates, P., O’Brien, S. J., Murphy, W. J., 2005. A molecular phylogeny for bats illuminates biogeography and the fossil record. Science 307, 580-584.
 
Tejedor, M. F., Czaplewski, N. J., Goin, F. J., Aragon, E., 2005. The oldest record of South American bats. Journal of Vertebrate Paleontology 25, 990-993.
 
Tong, Y.-S., 1997. Middle Eocene small mammals from Liguanqiao Basin of Henan province and Yuanqu Basin of Shanxi province, Central China. Palaeontologica Sinica 18, New Series C, 26, 1-256.
 
Veilleux, J. P., Whitaker, J. O., Veilleux, S. L., 2003. Tree-roosting ecology of reproductive female eastern pipistrelles, Pipistrellus subflavus, in Indiana. Journal of Mammalogy 84, 1068-1075.
 
Yu, W., Wu, Y., Yang, G., 2014. Early diversification trend and Asian origin for extent bat lineages. Journal of Evolutionary Biology 27, 2204-2218. http://dx.doi.org/10.1111/jeb.12477
 
Zachos, J., Pagani, M., Sloan, L., Thomas, E., Billups, K., 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292, 686-693.