Skip to main content

Deep time diversity of metatherian mammals: implications for evolutionary history and fossil-record quality

  • C. Verity Bennett (a1), Paul Upchurch (a2), Francisco J. Goin (a3) and Anjali Goswami (a4)

Despite a global fossil record, Metatheria are now largely restricted to Australasia and South America. Most metatherian paleodiversity studies to date are limited to particular subclades, time intervals, and/or regions, and few consider uneven sampling. Here, we present a comprehensive new data set on metatherian fossil occurrences (Barremian to end Pliocene). These data are analyzed using standard rarefaction and shareholder quorum subsampling (including a new protocol for handling Lagerstätte-like localities).

Global metatherian diversity was lowest during the Cretaceous, and increased sharply in the Paleocene, when the South American record begins. Global and South American diversity rose in the early Eocene then fell in the late Eocene, in contrast to the North American pattern. In the Oligocene, diversity declined in the Americas, but this was more than offset by Oligocene radiations in Australia. Diversity continued to decrease in Laurasia, with final representatives in North America (excluding the later entry of Didelphis virginiana) and Europe in the early Miocene, and Asia in the middle Miocene. Global metatherian diversity appears to have peaked in the early Miocene, especially in Australia. Following a trough in the late Miocene, the Pliocene saw another increase in global diversity. By this time, metatherian biogeographic distribution had essentially contracted to that of today.

Comparison of the raw and sampling-corrected diversity estimates, coupled with evaluation of “coverage” and number of prolific sites, demonstrates that the metatherian fossil record is spatially and temporally extremely patchy. Therefore, assessments of macroevolutionary patterns based on the raw fossil record (as in most previous studies) are inadvisable.

  • View HTML
    • Send article to Kindle

      To send this article to your Kindle, first ensure is added to your Approved Personal Document E-mail List under your Personal Document Settings on the Manage Your Content and Devices page of your Amazon account. Then enter the ‘name’ part of your Kindle email address below. Find out more about sending to your Kindle. Find out more about sending to your Kindle.

      Note you can select to send to either the or variations. ‘’ emails are free but can only be sent to your device when it is connected to wi-fi. ‘’ emails can be delivered even when you are not connected to wi-fi, but note that service fees apply.

      Find out more about the Kindle Personal Document Service.

      Deep time diversity of metatherian mammals: implications for evolutionary history and fossil-record quality
      Available formats
      Send article to Dropbox

      To send this article to your Dropbox account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Dropbox.

      Deep time diversity of metatherian mammals: implications for evolutionary history and fossil-record quality
      Available formats
      Send article to Google Drive

      To send this article to your Google Drive account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Google Drive.

      Deep time diversity of metatherian mammals: implications for evolutionary history and fossil-record quality
      Available formats
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (, which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.
Hide All
Abello M. A. 2013. Analysis of dental homologies and phylogeny of Paucituberculata (Mammalia: Marsupialia). Biological Journal of the Linnean Society 109:441465.
Alroy J. 1999. The fossil record of North American mammals: evidence for a Paleocene evolutionary radiation. Systematic Biology 48:107118.
Alroy J. 2008. Dynamics of origination and extinction in the marine fossil record. Proceedings of the National Academy of Sciences USA 105:1153611542.
Alroy J. 2010. Geographical, environmental and intrinsic biotic controls on Phanerozoic marine diversification. Palaeontology 53:12111235.
Alroy J. 2015. Current extinction rates of reptiles and amphibians. Proceedings of the National Academy of Sciences USA 112:1300313008.
Alroy J., Marshall C. R., Bambach R. K., Bezusko K., Foote M., Fürsich F. T., Hansen T. A., Holland S. M., Ivany L. C., Jablonski D., Jacobs D. K., Jones D. C., Kosnik M. A., Lidgard S., Low S., Miller A. I., Novack-Gottshall P. M., Olszewski T. D., Patzkowsky M. E., Raup D. M., Roy K., Sepkoski J. J., Sommers M. G., Wagner P. J., and Webber A.. 2001. Effects of sampling standardization on estimates of Phanerozoic marine diversification. Proceedings of the National Academy of Sciences USA 98:62616266.
Alroy J., Aberhan M., Bottjer D. J., Foote M., Fürsich F. T., Harries P. J., Hendy A. J., Holland S. M., Ivany L. C., Kiessling W., and Kosnik M. A.. 2008. Phanerozoic trends in the global diversity of marine invertebrates. Science 321:97100.
Antoine P. O., Marivaux L., Croft D. A., Billet G., Ganerød M., Jaramillo C., Martin T., Orliac M. J., Tejada J., Altamirano A. J., and Duranthon F.. 2012. Middle Eocene rodents from Peruvian Amazonia reveal the pattern and timing of caviomorph origins and biogeography. Proceedings of the Royal Society of Biology 279:13191326.
Archer M., Hand S., and Godthelp H.. 1988. A new order of Tertiary zalambdodont marsupials. Science 239:15281531.
Archibald J. D. 1983. Structure of the K–T mammal radiation in North America: speculations on turnover rates and trophic structure. Acta Palaeontologica Polonica 28:717.
Archibald J. D. 1996. Dinosaur extinction and the end of an era. Columbia University Press, New York.
Archibald J. D., and Bryant L. J.. 1990. Differential Cretaceous Tertiary extinctions of non-marine vertebrates—evidence from northeastern Montana. Geological Society of America Special Paper 247:549562.
Averianov A. O., Archibald J. D., and Ekdale E. G.. 2010. New material of the Late Cretaceous Deltatheroidan mammal Sulestes from Uzbekistan and phylogenetic reassessment of the metatherian-eutherian dichotomy. Journal of Systematic Palaeontology 8:301330.
Bacon C. D., Silvestro D., Jaramillo C., Smith B. T., Chakrabarty P., and Antonelli A.. 2015. Biological evidence supports an early and complex emergence of the Isthmus of Panama. Proceedings of the National Academy of Sciences USA 112:61106115.
Badgley C. 2003. The multiple scales of biodiversity. Paleobiology 29:1113.
Beck R. M. D. 2009. Was the Oligo-Miocene Australian metatherian Yalkaparidon a “mammalian woodpecker”? Biological Journal of the Linnean Society 97:117.
Beck R. M. D. 2012. An “ameridelphian” marsupial from the early Eocene of Australia supports a complex model of Southern Hemisphere marsupial biogeography. Naturwissenschaften 99:715729.
Beck R. M. D. 2015. A peculiar faunivorous metatherian from the early Eocene of Australia. Acta Palaeontologica Polonica 60:123129.
Bennett C. V., and Goswami A.. 2013. Statistical support for the hypothesis of developmental constraint in marsupial skull evolution. BMC Biology 11:52.
Benson R. B. J., and Butler R. J.. 2011. Uncovering the diversification history of marine tetrapods: ecology influences the effect of geological sampling biases. Geological Society of London Special Publication 358:191207.
Benson R. B. J., Butler R. J., Lindgren J., and Smith A. S.. 2010. Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates. Proceedings of the Royal Society of London B 277:829834.
Benton M. J. 2009. The Red Queen and court jester: species diversity and the role of biotic and abiotic factors through time. Science 323:728.
Benton M. J., Dunhill A. M., Lloyd G. T., and Marx F. G.. 2011. Assessing the quality of the fossil record: insights from vertebrates. Geological Society of London, Special Publication 358:6394.
Black K. H., Archer M., Hand S. J., and Godthelp H.. 2012. The rise of Australian marsupials: a synopsis of biostratigraphic, phylogenetic, palaeoecologic and palaeobiogeographic understanding. Pp 9831078 in J. A. Talent, ed. Earth and life. Springer, Dordrecht, Netherlands.
Black K. H., Price G. J., Archer M., and Hand S. J.. 2014. Bearing up well? Understanding the past, present and future of Australia’s koalas. Gondwana Research 25:11861201.
Blake D. B., and Aronson R. B.. 1998. Eocene stelleroids (Echinodermata) at Seymour Island, Antarctic Peninsula. Journal of Paleontology 72:339353.
Bohaty S. M., and Zachos J. C.. 2003. Significant Southern Ocean warming event in the late middle Eocene. Geology 31:10171020.
Bohaty S. M., Zachos J. C., Florindo F., and Delaney M. L.. 2009. Coupled greenhouse warming and deep‐sea acidification in the middle Eocene. Paleoceanography 24:PA2207.
Bond M., Tejedor M. F., Campbell K. E. Jr, Chornogubsky L., Novo N., and Goin F.. 2015. Eocene primates of South America and the African origins of New World monkeys. Nature 520:538541.
Bowen G. J., Maibauer B. J., Kraus M. J., Rohl U., Westerhold T., Steimke A., Gingerich P. D., Wing S. L., and Clyde W. C.. 2015. Two massive, rapid releases of carbon during the onset of the Palaeocene–Eocene thermal maximum. Nature Geoscience 8:4447.
Brusatte S., Butler R. J., Barrett P. M., Carrano M. T., Evans D. C., Lloyd G. T., Mannion P. D., Norell M. A., Pepp D. J., Upchurch P., and Williamson T. E.. 2015. The extinction of the dinosaurs. Biological Reviews 90:628642.
Bush A. M., Markey M. J., and Marshall C. R.. 2004. Removing bias from diversity curves: the effects of spatially organized biodiversity on sampling-standardization. Paleobiology 30:666686.
Butler R. J., Barrett P. M., Nowbath S., and Upchurch P.. 2009. Estimating the effects of sampling biases on pterosaur diversity patterns: implications for hypotheses of bird/pterosaur competitive replacement. Paleobiology 35:432446.
Butler R. J., Benson R. B. J., Carrano M. T., Mannnion P. D., and Upchurch P.. 2011. Sea level, dinosaur diversity and sampling biases: investigating the “common cause” hypothesis in the terrestrial realm. Proceedings of the Royal Society of London B 278:11651170.
Case J. A., Goin F. J., and Woodburne M .O.. 2004. “South American” marsupials from the Late Cretaceous of North America and the origin of marsupial cohorts. Journal of Mammalian Evolution 11:223255.
Cifelli R. L. 2004. Chapter 5: Marsupial mammals from the Albian–Cenomanian (Early–Late Cretaceous) boundary, Utah. Bulletin of the American Museum of Natural History 285:6279.
Cifelli R. L., and Davis B. M.. 2003. Paleontology. Marsupial origins. Science 302:18991900.
Cifelli R. L., Wible J. R., and Jenkins F. A.. 1998. Triconodont mammals from the Cloverly Formation (Lower Cretaceous), Montana and Wyoming. Journal of Vertebrate Paleontology 18:237241.
Cifelli R. L., Davis B. M., and Sames B.. 2014. Earliest Cretaceous mammals from the western United States. Acta Palaeontologica Polonica 59:3152.
Clemens W. A. 1968. Origin and early evolution of marsupials. Evolution 22:118.
Crampton J. S., Beu A. G., Cooper R. A., Jones C. M., Marshall B., and Maxwell P. A.. 2003. Estimating the rock volume bias in paleobiodiversity studies. Science 301:358360.
Davis B. M., Cifelli R. L., and Kielan-Jaworowska Z.. 2008. Earliest evidence of Deltatheroida (Mammalia: Metatheria) from the Early Cretaceous of North America. Pp. 324 in E. J. Sargis and M. Dagosto, eds. Mammalian evolutionary morphology: a tribute to Frederick S. Szalay. Springer, Dordrecht, Netherlands.
de la Fuente M. S., Salgado L., Albino A., Báez A. M., Bonaparte J. F., Calvo J. O., Chiappe L. M., Codorniú L. S., Coria R. A., Gasparini Z., and Riga B. J. González. 2007. Tetrápodos continentales del Cretácico de la Argentina: una síntesis actualizada. In Asociación Paleontológica Argentina Publicación Especial 11(Ameghiniana 50º aniversario): 137153. Buenos Aires, 25 November 2007.
de Muizon C., and Cifelli R. L.. 2000. The “condylarths” (archaic Ungulata, Mammalia) from the early Palaeocene of Tiupampa (Bolivia): implications on the origin of the South American ungulates. Geodiversitas 22:47150.
de Muizon C., and Cifelli R. L.. 2001. A new basal “didelphoid” (Marsupialia, Mammalia) from the early Paleocene of Tiupampa (Bolivia). Journal of Vertebrate Paleontology 21:8797.
dos Reis M., Inoue J., Hasegawa M., Asher R. J., Donoghue P. C., and Yang Z.. 2012. Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny. Proceedings of the Royal Society of London B 279:34913500.
dos Reis M., Donoghue P. C. J., and Yang Z.. 2014. Neither phylogenomic nor palaeontological data support a Palaeogene origin of placental mammals. Biology Letters 10:20131003.
Dowsett H. J., Barron J. A., Poore R. Z., Thompson R. S., Cronin T. M., Ishman S. E., and Willard D. A.. 1999 Middle Pliocene paleoenvironmental reconstruction: PRISM2. US Geological Survey Open File Report 99-535.
Echeverri S., Cardona A., Pardo A., Monsalve G., Valencia V. A., Borrero C., Rosero S., and Lopez S.. 2015. Regional provenance from southwestern Colombia fore-arc and intra-arc basins: implications for Middle to Late Miocene orogeny in the Northern Andes. Terra Nova 27:356363.
Ezcurra M. D., and Agnolin F. L.. 2012. A new global palaeobiogeographical model for the late Mesozoic and early Tertiary. Systematic Biology 61:553566.
Fastovsky D. E., Huang Y., Hsu J., Martin-McNaughton J., Sheehan P. M., and Weishampel D. B.. 2004. Shape of Mesozoic dinosaur richness. Geology 32:877880.
Flannery T. F, Turnbull W. D., Rich T. H. V., and Lundelius E. L.. 1987. The phalangerids (Marsupialia: Phalangeridae) of the early Pliocene Hamilton Local Fauna, Southwestern Victoria. Pp. 537546 in M. Archer, ed. Possums and opossums: studies in evolution. Surrey Beatty and Sons and the Royal Zoological Society of New South Wales, Sydney.
Flynn J. J., and Wyss A.. 1998. Recent advances in South American mammalian paleontology. Trends in Ecology and Evolution 13:449454.
Foote M. 2001. Inferring temporal patterns of preservation, origination, and extinction from taxonomic survivorship analysis. Paleobiology 27:602630.
Fröbisch J. 2008. Global taxonomic diversity of anomodonts (Tetrapoda, Therapsida) and the terrestrial rock record across the Permian-Triassic boundary. PLoS ONE 3: p. e3733.
Gaetano L. C., Marsicano C. A., and Rougier G. W.. 2013. A revision of the putative Late Cretaceous triconodonts from South America. Cretaceous Research 46:90100.
Gallagher S. J., Greenwood D. R., Taylor D., Smith A. J., Wallace M. W., and Holdgate G. R.. 2003. The Pliocene climatic and environmental evolution of southeastern Australia: evidence from the marine and terrestrial realm. Palaeogeography, Palaeoclimatology, Palaeoecology 193:349382.
Gayet M., Rage J. C., Sempere T., and Gagnier P. Y.. 1992. Modalités des échanges de vertébrés continentaux entre l’Amérique du Nord et l’Amérique du Sud au Crétacé supérieur et au Paléocène. Bulletin de la Société Géologique de France 163:781791.
Geoscience Australia and Australian Stratigraphy Commission. 2013. Australian Stratigraphic Units Database., accessed 1 May 2013.
Godthelp H., Archer M., and Cifelli R.. 1992. Earliest known Australian Tertiary mammal fauna. Nature 356:514516.
Goin F. J., Candela A. M., Abello M. A., and Oliveira E. V.. 2009. Earliest South American paucituberculatans and their significance in the understanding of “pseudodiprotodont” marsupial radiations. Zoological Journal of the Linnean Society 155:867884.
Goin F. J., Woodburne M. O., Zimicz A. N., Martin G. M., and Chornogubsky L.. 2016. A brief history of South American metatherians. Springer, Dordrecht, Netherlands.
Goin F. J., Woodburne M. O., Zimicz A. N., Martin G. M., and Chornogubsky L.. 2016. Paleobiology and adaptations of Paleogene metatherians. Pp. 185208 in A brief history of South American metatherians. Springer, Dordrecht, Netherlands.
Good I. J. 1953. The population frequencies of species and the estimation of population parameters. Biometrika 40:237264.
Goswami A. 2012. A dating success story: genomes and fossils converge on placental mammal origins. EvoDevo 3:18.
Goswami A., Randau M., Polly P. D., Weisbecker V., Bennett C. V., Hautier L., and Sánchez-Villagra M. R.. 2016. Do developmental constraints and high integration limit the evolution of the marsupial oral apparatus? Integrated Comparative Biology 56:404415.
Graham A. 2000. History of North American vegetation—Cretaceous (Maastrichtian)—Tertiary. Pp. 5770 in F.o.N.A.E. Committee ed. Flora of North America north of Mexico. Oxford University Press, New York.
Greenwood D., and Christophel D.. 2005. The origins and Tertiary history of Australian “tropical” rainforests. Pp. 336373 in E. Bermingham, C. Dick, and C. Moritz, eds. Tropical rainforests: past, present, future. University of Chicago Press, Chicago.
Grossnickle D. M., and Newham E.. 2016. Therian mammals experience an ecomorphological radiation during the Late Cretaceous and selective extinction at the K–Pg boundary. Proceedings of the Royal Society of London B 283:20160256.
Grossnickle D. M., and Polly P. D.. 2013. Mammal disparity decreases during the Cretaceous angiosperm radiation. Proceedings of the Royal Society of London B 280 280:20132110.
Halliday T. J. D., and Goswami A.. 2016. Eutherian morphological disparity across the end-Cretaceous mass extinction. Biological Journal of the Linnean Society 118:152168.
Halliday T. J. D., Upchurch P., and Goswami A.. 2016. Eutherians experience elevated evolutionary rates in the immediate aftermath of the Cretaceous–Palaeogene mass extinction. Proceedings of the Royal Society of London B 283:20153026.
Halliday T. J. D., Upchurch P., and Goswami A.. 2017. Resolving the relationship of Palaeocene placental mammals. Biological Reviews 92:521550.
Haq B. U., Hardenbol J., and Vail P. R.. 1987. Chronology of fluctuating sea levels since the Triassic. Science 235:11561167.
Harrell F. E., and Dupont C.. 2013. Hmisc: Harrell miscellaneous. R package, Version 3.10-1.1.
Hedges S. B. 2006. Paleogeography of the Antilles and the origin of West Indian terrestrial vertebrates. Annals of the Missouri Botanical Garden 93:231244.
Hill R. 1992. Nothofagus: evolution from a southern perspective. Trends in Ecology and Evolution 7:190194.
Hocknull S. 2005. Additional specimens of Bohra (Marsupialia: Macropodidae) from the Pliocene of Queensland. Memoirs of the Queensland Museum 51:26.
Hooker J. J. 2000. Palaeogene mammals: crises and ecological change. Pp. 333349 in S. Culver and P. Rawson, eds. Biotic response to global change: the last 145 million years. Cambridge University Press, Cambridge.
Hoorn C., and Flantua S.. 2015. An early start for the Panama land bridge. Science 348:186187.
Horovitz I., Martin T., Bloch J., Ladevèze S., Kurz C., and Sánchez-Villagra M. R.. 2009. Cranial anatomy of the earliest marsupials and the origin of opossums. PLoS ONE. doi: 10.1371/journal.pone.0008278.
Hu X., Garzanti E., Wang J., Huang W., An W., and Webb A.. 2016. The timing of India-Asia collision onset—facts, theories, controversies. Earth Science Reviews 160:264299.
Hunt R., and Poole I.. 2003. Paleogene West Antarctic climate and vegetation history in light of new data from King George Island. In S. L. Wing, P. Gingerich, B. Schmitz, and E. Thomas, eds. Causes and consequences of globally warm climates in the early Paleogene. Geological Society of America Special Paper 369:395412.
Huyghe D., Lartaud F., Emmanuel L., Merle D., and Renaud M.. 2015. Palaeogene climate evolution in the Paris Basin from oxygen stable isotope (δ18O) compositions of marine molluscs. Journal of the Geological Society 172:576587.
Iturralde-Vinent M., and MacPhee R. D. E.. 1999. Paleogeography of the Caribbean region: implications for Cenozoic biogeography. Bulletin of the American Museum of Natural History 238:195.
Ivany L. C., Lohmann K. C., Hasiuk F., Blacke D. B., Glass A., Aronson R. B., and Moody R. M.. 2008. Eocene climate record of a high southern latitude continental shelf: Seymour Island, Antarctica. Geological Society of America Bulletin 120:659678.
Jablonski D., Roy K., Valentine J. W., Price R. M., and Anderson P. S.. 2003. The impact of the pull of the recent on the history of marine diversity. Science 360:11331135.
Janis C. M. 1993. Tertiary mammal evolution in the context of changing climates, vegetation, and tectonic events. Annual Review of Ecology and Systematics 24:467500.
Jansa S. A., Barker F. K., and Voss R. S.. 2014. The early diversification history of didelphid marsupials: a window into South America’s “Splendid Isolation.” Evolution 68:684695.
Jaramillo C. 2002. Response of tropical vegetation to Paleogene warming. Paleobiology 28:222243.
Jovane L., Florindo F., Coccioni R., Dinarès-Turell J., Marsili A., Monechi S., Roberts A. P., and Sprovieri M.. 2007. The middle Eocene climatic optimum event in the Contessa Highway section, Umbrian Appennines, Italy. Geological Society of America Bulletin 119:413427.
Kalmar A., and Currie D. J.. 2010. The completeness of the continental fossil record and its impact on patterns of diversification. Paleobiology 36:5160.
Kelly E. M., and Sears K. E.. 2011. Limb specialization in living marsupial and eutherian mammals: constraints on mammalian limb evolution. Journal of Mammalogy 92:10381049.
Kennett J. 1977. Cenozoic evolution of Antarctic glaciation, the Circum-Antarctic Ocean, and their impact on global paleoceanography. Journal of Geophysical Research 82:38433860.
Kennett J. P. 1985. Neogene paleoceanography and plankton evolution. South African Journal of Science 81:251253.
Kielan-Jaworowska Z., and Cifelli R. L.. 2001. Primitive boreosphenidan mammal (?Deltatheroida) from the Early Cretaceous of Oklahoma. Acta Palaeontologica Polonica 46:377391.
Kielan-Jaworowska Z., and Nessov L. A.. 1990. On the metatherian nature of the Deltatheroida, a sister group of the Marsupialia. Lethaia. 23:110.
Kielan-Jaworowska Z., Cifelli R. L., and Luo Z.-X.. 2004. Mammals from the age of dinosaurs: structure, relationships, and paleobiology. Columbia University Press, New York.
Kohn M. J., Stromberg C. A. E., Madden R. H., Dunn R. E., Evans S., Palacios A., and Carlini A. A.. 2015. Quasi-static Eocene–Oligocene climate in Patagonia promotes slow faunal evolution and mid-Cenozoic global cooling. Palaeogeography, Palaeoclimatology and Palaeoecology 435:2437.
Krause D. W., Hoffmann S., Wible J. R., Kirk E. C., Schultz J. A., von Koenigswald W., Groenke J. R., Rossie J. B., O’Connor P. M., Seiffert E. R., and Dumont E. R.. 2014. First cranial remains of a gondwanatherian mammal reveal remarkable mosaicism. Nature 515:512517.
Krockenberger A. 2006. Lactation. Pp. 108136 in P. Armati, C. Dickman, and I. Hume, eds. Marsupials. Cambridge University Press, Cambridge.
La Roux J. P. 2012. A review of Tertiary climate changes in southern South America and the Antarctic Peninsula. Part 2: continental conditions. Sedimentary Geology 247–248:2138.
Lawver L., and Gahagan L.. 2003. Evolution of Cenozoic seaways in the circum-Antarctic region. Palaeogeography, Palaeoclimatology, Palaeoecology 198:1137.
Legendre S. 1982. La faune de micromammiferes du gisement burdigalien de Port-la-Nouvelle (Aude); complements et indications paleoecologiques. Bulletin de la Societe geologique de France 24:383387.
Lillegraven J. A. 1974. Biogeographical considerations of the marsupial-placental dichotomy. Annual Review of Ecology and Systematics 5:263283.
Lillegraven J. A. 1975. Biological considerations of the marsupial-placental dichotomy. Evolution 29:707722.
Liow L. H., Fortelius M., Bingham E., Lintulaakso K., Mannila H., Flynn L., and Stenseth N. S.. 2008. Higher origination and extinction rates in larger mammals. Proceedings of the National Academy of Sciences USA 105:60976102.
Lisiecki L. E., and Raymo M. E.. 2007. Plio–Pleistocene climate evolution: trends and transitions in glacial cycle dynamics. Quaternary Science Reviews 26:5669.
Lloyd G. T., and Friedman M.. 2013. A survey of palaeontological sampling biases in fishes based on the Phanerozoic record of Great Britain. Palaeogeography, Palaeoclimatology, Palaeoecology 372:517.
Longrich N. R., Scriberas J., and Wills M. A.. 2016. Severe extinction and rapid recovery of mammals across the Cretaceous–Paleogene boundary, and the effects of rarity on patterns of extinction and recovery. Journal of Evolutionary Biology 29:14951512.
Lowen M. A., Irmins R. B., Sertich J. J. W., Currie P. J., and Sampson S. D.. 2013. Tyrant dinosaur evolution tracks the rise and fall of Late Cretaceous oceans. PLoS ONE 8:e79420.
Luo Z.-X., Cifelli R. L., and Kielan-Jaworowska Z.. 2001. Dual origin of tribosphenic mammals. Nature 409:5357.
Luo Z.-X., Ji Q., Wible J. R., and Yuan C.-X.. 2003. An Early Cretaceous tribosphenic mammal and metatherian evolution. Science 302:19341940.
Luo Z.-X., Yuan C.-X., Meng Q. J., and Ji Q.. 2011. A Jurassic eutherian mammal and divergence of marsupials and placentals. Nature 476:442445.
Mannion P. D., Upchurch P., Carrano M. T., and Barrett P. M. 2011. Testing the effect of the rock record on diversity: a multidisciplinary approach to elucidating the generic richness of sauropodomorph dinosaurs through time. Biological Reviews of the Cambridge Philosophical Society 86:157181.
Mannion P. D., Benson R. B. J., Carrano M. T., Tennant J. P., Judd J., and Butler R. J.. 2015. Climate constrains the evolutionary history and biodiversity of crocodylians. Nature Communications 6:19.
Marshall L. G. 1980. Systematics of the South American marsupial family Caenolestidae. Fieldiana 5:1145.
Martin H. 1994. Australian Tertiary phytogeography. Pp. 104142 in R. Hill, ed. History of the Australian vegetation. Cambridge University Press, Cambridge.
Martin H. 2006. Cenozoic climatic change and the development of the arid vegetation in Australia. Journal of Arid Environments 66:533563.
McGowran B., and Hill R. S.. 2015. Cenozoic climatic shifts in southern Australia. Transactions of the Royal Society of South Australia 139:1937.
McGowran B., Archer M., Bock P., Sarragh T. A., Godthelp H., Hageman S., Hand S. J., Hill R., Li Q., Maxwell P. A., McNamara K. J., Macphail M., Mildenhall D., Partridge A. D., Richardson J., Shafik S., Truswell E. M., and Warne M.. 2000. Australasian palaeobiogeography: the Palaeogene and Neogene record. Memoirs of the Association of Australasian Palaeontologists. 23:405470.
McNab B. 1986. Food habits, energetics and the reproduction of marsupials. Journal of Zoology 208:595614.
McNab B. 2005. Uniformity in the basal metabolic rate of marsupials: its causes and consequences. Revista Chilena De Historia Natural 78:183198.
McNab B. 2008. An analysis of the factors that influence the level and scaling of mammalian BMR. Comparative Biochemistry and Physiology A 151:528.
Mehrotra R. 2003. Status of plant megafossils during the early Paleogene in India. In S. L. Wing, P. Gingerich, B. Schmitz, and E. Thomas, eds. Causes and consequences of globally warm climates in the early Paleogene. Geological Society of America Special Paper 369:413423.
Meredith R. W., Janečka J. E., Gatesy J., Ryder O. A., Fisher C. A., Teeling E. C., Goodbla A., Eizirik E., Simão T. L., Stadler T., and Rabosky D. L.. 2011. Impacts of the Cretaceous terrestrial revolution and KPg extinction on mammal diversification. Science 334:521524.
Miller A. I., and Foote M.. 1996. Calibrating the Ordovician radiation of marine life: implications for Phanerozoic diversity trends. Paleobiology 22:304309.
Molnar P. 2008. Closing of the Central American Seaway and the Ice Age: a critical review. Paleoceanography 23:PA2201.
Montes C., Cardona A., Jaramillo C., Pardo A., Silva J. C., Valencia V., Ayala C., Pérez-Angel L. C., Rodriguez-Parra L. A., Ramirez V., and Niño H.. 2015. Middle Miocene closure of the Central American seaway. Science 348:226229.
Montes M., Nozal F., Santillana S., Marenssi S., and Olivero E.. 2013. Mapa Geologico de la isla Marambio (Seymour) Escala 1:20.000 Primera Edicion. Serie Cartografica Geocientıfica Antartica. Madrid-Instituto Geologico y Minero de Espana. Instituto Antartico Argentino, Buenos Aires.
Moran K., Backman J., Brinkhuis H., Clemens S. C., Cronin T., Dickens G. R., Eynaud F., Gattacceca J., Jakobsson M., Jordan R. W., Kaminski M., King J., Koc N., Krylov A., Martinez N., Matthiessen J., McInroy D., Moore T. C., Onodera J., O’Regan M., Pälike H., Rea B., Rio D., Sakamoto T., Smith D. C., Stein R., St John K., Suto I., Suzuki N., Takahashi K., Watanabe M., Yamamoto M., Farrell J., Frank M., Kubik P., Jokat W., and Kristoffersen Y.. 2006. The Cenozoic palaeoenvironment of the Arctic Ocean. Nature 441:601605.
Morley R. J. 2000. The origin and evolution of tropical rain forests. Wiley, New York.
Mörner N. A., and Sylwan C.. 1989. Magnetostratigraphy of the Patagonian moraine sequence at Lago Buenos Aires. Journal of South American Earth Sciences 2:385389.
Morton S. R., Recher H. F., Thompson S. D., and Braithwaite R. W.. 1982. Comments on the relative advantages of marsupial and eutherian reproduction. American Naturalist 120:128134.
Muirhead J., Dawson L., and Archer M.. 1997. Perameles bowensis, a new species of Perameles (Peramelemorphia, Marsupialia) from Pliocene faunas of Bow and Wellington Caves, New South Wales. Proceedings of the Linnean Society of New South Wales 117:163174.
Nessov L. A. 1985. Redkie kostnye ryby, nazemiye yashcheritsy i mlekopitayushchie zony limanov i primorskikh nizmennostey mela Kyzylkumov [Rare bony fishes, terrestrial lizards, and mammals in the estuarine and coastal lowland zone of the Kyzyl-Kum Cretaceous]. Yearbook of the All-Union Paleontological Association 28:199219.
Newham E., Benson R. B. J., Upchurch P., and Goswami A.. 2014. Mesozoic mammaliaform diversity: the effect of sampling corrections on reconstructions of evolutionary dynamics. Palaeogeography, Palaeoclimatology, Palaeoecology 412:3244.
Nicholson D. B., Holroyd P. A., Benson R. B. J., and Barrett P. M.. 2015. Climate-mediated diversification of turtles in the Cretaceous. Nature Communications 6:18.
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., and Cirranello A. L.. 2013. The placental mammal ancestor and the post–K–Pg radiation of placentals. Science 339:662667.
Ortiz-Jaureguizar E. 2003. Relaciones de similitud, paleoecología y extinción de los Abderitidae (Marsupialia, Paucituberculata). Coloquios de Paleontología, Extra Volume 1:475498.
Ortiz-Jaureguizar E., and Cladera G. A.. 2006. Paleoenvironmental evolution of southern South America during the Cenozoic. Journal of Arid Environments 66:498532.
Ortiz-Jaureguizar E., and Pascual R.. 2007. The tectonic setting of the Caribbean region: key in the radical Late Cretaceous–Early Paleocene South American land-mammal turnover. Pp. 301307 in E. Diaz-Martinez and I. Rabano, eds. Fourth European Meeting on the Palaeontology and Stratigraphy of Latin America. Instituto Geologico y Minero de Espana, Madrid.
Owen R. 1838. Appendix to Mitchell, T. L. Fossil remains from Wellington Valley, Australia. Marsupialia. Pp. 359369 in Three expeditions into the interior of Eastern Australia, with descriptions of the recently explored region of Australia Felix, and of the present colony of New South Wales. T. and W, Boone, London.
Padian K., and Clemens W.. 1985. Terrestrial vertebrate diversity: episodes and insights. Pp. 4196 in J. W. Valentine, ed. Phanerozoic diversity patterns: profiles in macroevolution. Princeton University Press, Princeton, N.J.
Palazzesi L., and Barreda V.. 2012. Fossil pollen records reveal a late rise of open-habitat ecosystems in Patagonia. Nature Communications 3:1294.
Palazzesi L., Barreda V. D., Cuitino J. I., Guler M. V., Telleria M. C., and Santos R. V.. 2014. Fossil pollen records indicate that Patagonian desertification was not solely a consequence of Andean uplift. Nature Communications 5:3558.
Pascual R., and Ortiz-Jaureguizar E.. 2007. The Gondwanan and South American episodes: two major and unrelated moments in the history of the South American mammals. Journal of Mammalian Evolution 14:75137.
Pascual R., Ortiz-Jaureguizar E., and Prado J. L.. 1996. Land mammals: paradigm of Cenozoic South American geobiotic evolution. In. Contribution of Southern South America to Vertebrate Paleontology. Muncher Geowissenschaftliche Abhandlungen (A) 30:265319.
Patterson B., and Pascual R.. 1972. The fossil mammal fauna of South America. Pp. 247309 in A. Keast, F. C. Erk, and B. Glass, eds. Evolution, mammals, and southern continents. State University of New York Press, Albany.
Pearson M. R., Benson R. B. J., Upchurch P., Fröbisch J., and Kammerer C. F.. 2013. Reconstructing the diversity of early terrestrial herbivorous tetrapods. Palaeogeography, Palaeoclimatology, Palaeoecology 372:4249.
Peters S. E., and Foote M.. 2001. Biodiversity in the Phanerozoic: a reinterpretation. Paleobiology 27:583601.
Pledge N. S. 1987. Phascolarctos maris, a new species of koala (Marsupialia: Phascolarctidae) from the Early Pliocene of South Australia. Pp. 327330 in M. Archer, ed. Possums and opossums: studies in evolution. Surrey Beatty and Sons and the Royal Zoological Society of New South Wales, Sydney.
Pough F. H., Andrews R. M., Cadle J. E., Crump M. L., Savitzky A. H., and Wells K. D.. 2004. Herpetology, 3rd ed. Benjamin Cummings, Upper Saddle River, N.J.
Prevosti F. J., Forasiepi A., and Zimicz N.. 2013. The evolution of the Cenozoic terrestrial mammalian predator guild in South America: competition or replacement? Journal of Mammalian Evolution 20:321.
Quattrocchio M., Durango de Cabrera J., and Galli C.. 2003. Formación Anta (Mioceno Temprano/Medio), Subgrupo Metán (Grupo Orán), en el río Piedras, Pcia. de Salta. Datos palinológicos. Revista de la Asociación Geológica Argentina 58:117127.
R Development Core Team. 2016. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
Rage J.-C. 1981. Les continents péri-atlantiques au Crétacé Supérieur: Migrations des faunes continentales et problèmes paléogéographiques. Cretaceous Research 2:6584.
Rage J.-C. 1986. South American/North American terrestrial interchanges in the latest Cretaceous: short comments on Brett-Surman and Paul (1985), with additional data. Journal of Vertebrate Paleontology 6:382383.
Rana R. S., and Wilson G. P.. 2003. New Late Cretaceous mammals from the Intertrappean beds of Rangapur, India and paleobiogeographic framework. Acta Palaeontologica Polonica 48:331348.
Raup D. M. 1972. Taxonomic diversity during the Phanerozoic. Science 177:10651071.
Raup D. M. 1975. Taxonomic diversity estimation using rarefaction. Paleobiology 1:333342.
Raup D. M. 1979. Biases in the fossil record of species and genera. Bulletin of the Carnegie Museum of Natural History 13:8591.
Rea D. K., Zachos J. C., Owen R. M., and Gingerich P. D.. 1990. Global change at the Paleocene–Eocene boundary: climatic and evolutionary consequences of tectonic events. Palaeogeography, Palaeoclimatology, Palaeoecology 79:117128.
Reeder D. M., Helgen K. M., and Wilson D. E.. 2007. Global trends and biases in new mammal species discoveries. Museum of Texas Tech University Occasional Paper 269.
Riggs E. S. 1934. A new marsupial saber-tooth from the Pliocene of Argentina and its relationships to other South American predacious marsupials. Transactions of the American Philosophical Society 24:132.
Rougier G. W., Wible J. R., and Novacek M. J.. 1998. Implications of Deltatheridium specimens for early marsupial history. Nature 396:459463.
Sahney S., Benton M. J., and Ferry P.. 2010. Links between global taxonomic diversity, ecological diversity and the expansion of vertebrates on land. Biology Letters 6:544547.
Sánchez-Villagra M. R. 2013. Why are there fewer marsupials than placentals? On the relevance of geography and physiology to evolutionary patterns of mammalian diversity and disparity. Journal of Mammalian Evolution 20:279290.
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., and Collins G. S.. 2010. The Chicxulub asteroid impact and mass extinction at the Cretaceous–Paleogene boundary. Science 327:2141218.
Sears K. E. 2004. Constraints on the morphological evolution of marsupial shoulder girdles. Evolution 58:23532370.
Seton M., Müller R. D., Zahirovic S., Gaina C., Torsvik T., Shepard G., Talsma A., Gurnis M., Turner M., Maus S., and Chandler M.. 2012. Global continental and ocean basin reconstructions since 200 Ma. Earth-Science Reviews 113:212270.
Shackleton N. J. 1995. New data on the evolution of Pliocene climate variability. Pp. 242248 in E. S. Vrba, ed. Paleoclimate and evolution, with emphasis on human origins. Yale University Press, New Haven, Conn.
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., and Westberg-Smith J.. 1984. Oxygen isotope calibration of the onset of ice-rafting and history of glaciation in the North Atlantic region. Nature 307:620623.
Shellito C. J., and Sloan L. C.. 2006. Reconstructing a lost Eocene paradise. Part I: simulating the change in global floral distribution at the initial Eocene thermal maximum. Global and Planetary Change 50:117.
Signor P. W. I., and Lipps J. H.. 1982. Sampling bias, gradual extinction patterns and catastrophes in the fossil record. Geological Society of America Special Paper 190:291296.
Simons E. L., and Bown T. M.. 1984. A new species of Peratherium (Didelphidae; Polyprotodonta): the first African marsupial. Journal of Mammalogy 65:539548.
Slater G. J. 2013. Phylogenetic evidence for a shift in the mode of mammalian body size evolution at the Cretaceous–Palaeogene boundary. Methods in Ecology and Evolution 4:734744.
Smith A. B., and McGowan A. J. M.. 2007. The shape of the Phanerozoic marine palaeodiversity curve: how much can be predicted from the sedimentary record of western Europe? Palaeontology 50:765774.
Smith A. S., and Benson R. B. J.. 2013. Marine diversity in the geological record and its relationship to surviving bedrock area, lithofacies diversity, and original marine shelf area. Geology 41:171174.
Tennant J. P., Mannion P. D., Upchurch P., Sutton M., and Price G.. 2017. Biotic and environmental dynamics across the Jurassic/Cretaceous boundary: evidence for a protracted period of faunal and ecological turnover. Biological Reviews 92:776814.
Travouillon K. J., Legendre S., Archer M., and Hand S. J.. 2009. Palaeoecological analyses of Riversleigh’s Oligo-Miocene sites: implications for Oligo-Miocene climate change in Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 276:2437.
Travouillon K. J., Gurovich Y., Beck R. M. D., and Muirhead J.. 2010. An exceptionally well-preserved short-snouted bandicoot (Marsupialia; Peramelemorphia) from Riversleigh’s Oligo-Miocene deposits, northwestern Queensland, Australia. Journal of Vertebrate Paleontology 30:15281546.
Trofimov B. A., and Szalay F. S.. 1994. New Cretaceous marsupial from Mongolia and the early radiation of Metatheria. Proceedings of the National Academy of Sciences USA 91:1256912573.
University of California Museum of Paleontology. 2016. Collections database., accessed 1 May 2016.
Uhen M. D., and Pyenson N. D.. 2007. Diversity estimates, biases and historiographic effects: resolving cetacean diversity in the Tertiary. Palaeontologica Electronica 10:11A–22.
Uliana M. A., and Biddle K. T.. 1988. Mesozoic–Cenozoic paleogeographic and geodynamic evolution of southern South America. Revista Brasileira de Geociencias 18:172190.
Upchurch P., Mannion P. D., Benson R. B. J., Butler R. J., and Carrano M. T.. 2011. Geological and anthropogenic controls on the sampling of the terrestrial fossil record: a case study from the Dinosauria. Geological Society of London Special Publication 358:209240.
U.S. Geological Survey. 2013. Home page., accessed 1 May 2013.
Vermeij G. J., and Leighton L. R.. 2003. Does global diversity mean anything? Paleobiology 29:37.
Villa G., Fioroni C., Pea L., Bohaty S., and Persico D.. 2008. Middle Eocene–late Oligocene climate variability: calcareous nannofossil response at Kerguelen Plateau, Site 748. Marine Micropaleontology 69:173192.
Vullo R., Gheerbrant E., de Muizon C., and Néraudeau D.. 2009. The oldest modern therian mammal from Europe and its bearing on stem marsupial paleobiogeography. Proceedings of the National Academy of Sciences USA 106:1991019915.
Wall P. D., Ivany L. C., and Wilkinson B. H.. 2009. Revisiting Raup: exploring the influence of outcrop area on diversity in light of modern sample-standardization techniques. Paleobiology 35:146167.
Whistler D. P., and Lander E. B.. 2003. Chapter 11: New late Uintan to early Hemingfordian land mammal assemblages from the undifferentiated Sespe and Vaqueros Formations, Orange County, and from the Sespe and Equivalent marine formations in Los Angeles, Santa Barbara, and Ventura Counties, Southern California. Bulletin of the American Museum of Natural History 279:231268.
Wible J. R., Rougier G. W., Novacek M. J., and Asher R. J.. 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary. Nature 447:10031006.
Wilf P. N., Cúneo R., Escapa I. H., Pol D., and Woodburne M. O.. 2013. Splendid and seldom isolated: the paleobiogeography of Patagonia. Annual Review of Earth and Planetary Sciences 41:561.
Williamson T. E., Brusatte S. L., Carr T. D., Weil A., and Standhardt B. R.. 2012. The phylogeny and evolution of Cretaceous–Palaeogene metatherians: cladistic analysis and description of new early Palaeocene specimens from the Nacimiento Formation, New Mexico. Journal of Systematic Palaeontology 10:625651.
Williamson T. E., Brusatte S. L., and Wilson G. P.. 2014. The origin and early evolution of metatherian mammals: the Cretaceous record. ZooKeys 465:176.
Wilson D. E., and Reeder D. M.. 2005. Mammal species of the world. Johns Hopkins University Press, Baltimore, Md.
Wilson G. P. 2013. Mammals across the K/Pg boundary in northeastern Montana, U.S.A.: Dental morphology and body size patterns reveal extinction selectivity and immigrant-fueled ecospace filling. Paleobiology 39:429469.
Wilson G. P. 2014. Mammalian extinction, survival, and recovery dynamics across the Cretaceous-Paleogene boundary in northeastern Montana, USA. Pp. 365392 in G. P. Wilson, W. A. Clemens, J. H. Horner, and J. H. Hartman, eds. Through the end of the Cretaceous in the type locality of the Hell Creek Formation in Montana and adjacent areas. Geological Society of America Special Paper 503.
Wilson G. P., Evans A. R., Corfe I. J., Smits P. D., Fortelius M., and Jernvall J.. 2012. Adaptive radiation of multituberculate mammals before the extinction of dinosaurs. Nature 483:457.
Wing S. L. 1998. Tertiary vegetation of North America as a context for mammalian evolution. In C. M. Janis, K. M. Scott, and L. L. Jacobs, eds. Evolution of Tertiary mammals of North America 1:3765. Cambridge University Press, Cambridge.
Wing S. L., Harrington G. J., Bloch J. I., Boyer D. M., and Smith F.. 2004. Major transient floral change during the Paleocene-Eocene thermal maximum. American Geophysical Union Fall Meeting, Abstracts, PP14A-07.
Wolfe J. A., and Dilcher D. L.. 1999. Late Paleocene through Middle Eocene Climates in Lowland North America. Pp. 184185 in F. P. Andreasson, B. Schmitz, and E. I. Thompson, eds. Early Paleogene warm climate and biosphere dynamics. Earth Sciences Center, Goteborg University, Goteborg, Sweden.
Woodburne M. O. 2004. Late Cretaceous and Cenozoic mammals of North America: biostratigraphy and geochronology. Columbia University Press, New York.
Woodburne M. O. 2010. The Great American Biotic Interchange: dispersals, tectonics, climate, sea level and holding pens. Journal of Mammalian Evolution 17:245264.
Zachos J., Pagani M., Sloan L., Thomas E., and Billups K.. 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292:686693.
Zhang P. Z., Molnar P., and Downs W. R.. 2001. Increased sedimentation rates and grain sizes 2-4 Myr ago due to the influence of climate change on erosion rates. Nature 410:891897.
Ziegler R. 1990. Didelphidae, Erinaceidae, Metacodontidae und Dimylidae (Mammalia) aus dem Oberoligozän und Untermiozän Süddeutschlands. Stuttgarter Beiträge zur Naturkunde, Serie B 158:199.
Zimicz N. 2014. Avoiding competition: the ecological history of late Cenozoic metatherian carnivores in South America. Journal of Mammalian Evolution 21:383393.
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

  • ISSN: 0094-8373
  • EISSN: 1938-5331
  • URL: /core/journals/paleobiology
Please enter your name
Please enter a valid email address
Who would you like to send this to? *


Altmetric attention score

Full text views

Total number of HTML views: 46
Total number of PDF views: 329 *
Loading metrics...

Abstract views

Total abstract views: 953 *
Loading metrics...

* Views captured on Cambridge Core between 6th February 2018 - 20th February 2018. This data will be updated every 24 hours.