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Division of Biogeoscience
Division of Geology and Geochemistry
Division of Urban Geograpahy and Regional Studies
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P.O. Box 64
(Gustaf Hällströmin katu 2a)
FI-00014 University of Helsinki

Department Office
Institute of Seismology

P.O. Box 68
(Gustaf Hällströmin katu 2b)
FI-00014 University of Helsinki

phone +358 294 1911 (university exchange)
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Mikael Fortelius, Ph.D.

Fortelius, photo by Daniel Burgas


Professor of Evolutionary Palaeontology
–paleontology and paleoecology

Department of Geosciences and Geography
P.O.Box 64 (Gustaf Hällströmin katu 2a)
FIN-00014 University of Helsinki

Tel: +358 2941 50837 or +358 50 533 7267

Research interests

  • Evolution of mammal communities and circumstances of early human environments
  • Evolution of Eurasian land mammals and terrestrial environments during the late Cenozoic (the last 24 million years)
  • Ecometrics of fossil and living mammals
  • Computational approaches to palaeobiology, especially analysis of temporal and spatial patterns
  • Developmental biology, function and evolution of mammalian teeth
  • Scaling problems (and solutions)

Teaching topics

  • Life and earth history
  • Development, morphology, function and evolution of teeth
  • Past terrestrial environments and ecosystems
  • Mammalian and human evolution
  • Basics of fossil recognition

Research Statement

I’m a palaeontologist with special interest in plant-eating mammals and their relationship with habitat and climate change. I’m particularly fascinated by mammalian teeth, how they form, how they work, how they wear down, and how their shapes evolve in evolutionary time. My first academic training was in biology, and my teacher in palaeontology was Björn Kurtén, who had a deeply organismic attitude to fossils. I carry with me a conviction that past organisms and ecosystems can only really be understood in relation to the living world. Even more strongly I feel that we cannot hope to understand the living world without sound and detailed knowledge of its stupendously long history. I’m somewhat passionate about resolving fossil data geographically and estimating the population sizes as well as the numbers of extinct species.

I started out as a lonesome specialist on fossil rhinoceroses and pigs, but most of my later research has been collaborative. It currently includes seven main areas: deep time palaeoecology and evolution of mammal communities, especially of Eurasia during the last 24 million years; ecometrics of mammalian teeth, especially palaeodiet reconstruction; multidisciplinary fieldwork, most recently in the late Miocene (12 to 5 million years ago) of North China; exploration of modern computational methods for the analysis of large palaeontological datasets (e.g., temporal seriation and analysis of spatial patterns); palaeoclimate reconstruction and modelling; environmental controls on the spatial distribution of modern species and communities; and scaling questions, especially of mammalian teeth and sense organs. I also participate in work on developmental biology and evolution of mammalian teeth. Lately I have become much involved with the fossil mammals and environments associated with early humans in East Africa, especially through collaboration with Meave Leakey and her research network.

Since 1992, I’ve been the coordinator of a public database of Neogene Old World Mammals. I’m also part of the MorphoBrowser project, a public database of 3D tooth shape.

I try to speak intelligently in public about evolution, including human origins, for example at the bi-annual Science Forum in Helsinki and recently on Darwin Day in Oslo. I’ve also participated in the production of educational programmes and documentaries, including Walking with Beasts of the BBC and Discovery Channel.

Among other things I’m also a Member of the Finnish Society of Sciences and Letters as well as the Finnish Academy of Science and Letters, and Editor (one of many) of Evolutionary Ecology Research.

Curriculum Vitae

Recent publications

Rannikko JC, Zliobaite I, Fortelius M. 2017. Relative abundances and palaeoecology of four suid genera in the Turkana Basin, Kenya, during the late Miocene to Pleistocene. Palaeogeography, Palaeoclimatology, Palaeoecology.

Zliobaite I, Puolamäki K, Eronen J, Fortelius M. 2017 A survey of computational methods for fossil data analysis. Evolutionary Ecology Research.

Patterson DB, Brown DB, Behrensmeyer AK, Merritt S, Zliobaite I, Reeves JS et al. 2017. Ecosystem evolution and hominin paleobiology at East Turkana, northern Kenya between 2.0 and 1.4 Ma. Palaeogeography, Palaeoclimatology, Palaeoecology 481:1-13. Available from, DOI: 10.1016/j.palaeo.2017.05.001

Barnosky, AD, Hadly, EA, Gonzalez, P, Head, J, Polly, JP, Lawing, AM, Eronen, JT, Ackerly, DD Ken Alex, Biber, E, Blois, J, Brashares, J, Ceballos, G, Davis, E, Dietl, GP, Dirzo, R, Doremus, H, Fortelius, M, Greene, HW, Hellmann, J, Hickler, T, Jackson, ST, Kemp, M, Koch, PL, Kremen, C, Lindsey, EL, Looy, C, Marshall, CR, Mendenhall, C, Mulch, A, Mychajliw, AM, Nowak, C, Ramakrishnan, U, Schnitzler, J, Das Shrestha, K, Solari, K, Stegner, L, Stegner, MA, Stenseth, NC, Wake, MH, Zhang, Z. 2017. Merging paleobiology with conservation biology to guide the future of terrestrial ecosystems. Science 355:eaah4787. doi: 10.1126/science.aah4787

Raia P, (ed.), Fortelius M, (ed.). 2017 Evolutionary macroecology. Evolutionary Ecology Research 18.

Raia P, Fortelius M. 2017. Introduction to Evolutionary Macroecology. Evolutionary Ecology Research 18:1-6.

Castiglione S, Mondanaro A, Carotenuto F, Passaro F, Fortelius M, Raia P. 2017. The many shapes of diversity: ecological and evolutionary determinants of biodiversity through time. Evolutionary Ecology Research 18:25-39.

Zliobaite, I., Rinne, J., Tóth, A., Mechenich, M, Liu, L., Behrensmeyer, A.K., and Fortelius, M. 2016. Herbivore teeth predict climatic limits in Kenyan ecosystems. PNAS 113: 12751-12756. doi: 10.1073/pnas.1609409113

Zliobaite I, Fortelius M. 2016. Revise rules on conflicts of interest. Nature 539(7628):168.

Mirzaie Ataabadi M, (ed.), Fortelius M, (ed.). 2016. The late Miocene Maragheh mammal fauna; results of recent multidisciplinary research. Palaeobiodiversity and Palaeoenvironments 96(3), 167 p.

Mirzaie Ataabadi M, Fortelius M. 2016. Introduction to the special issue "The late Miocene Maragheh mammal fauna; results of recent multidisciplinary research''. Palaeobiodiversity and Palaeoenvironments 96(3):339-347. Available from, DOI: 10.1007/s12549-016-0242-3

Sawada Y, Zaree GR, Sakai T, Itaya T, Yagi K, Imaizumi M et al. 2016. K-Ar ages and petrology of the late Miocene pumices from the Maragheh Formation, northwest Iran. Palaeobiodiversity and Palaeoenvironments 96(3):399-431. Available from, DOI: 10.1007/s12549-016-0232-5

Sakai T, Zaree G, Sawada Y, Mirzaie Ataabadi M, Fortelius M. 2016. Depositional environment reconstruction of the Maragheh Formation, East Azarbaijan, Northwestern Iran. Palaeobiodiversity and Palaeoenvironments. 96(3):383-398. Available from, DOI: 10.1007/s12549-016-0238-z

Karme A, Ungar PS, Healy C, Teaford M, Fortelius M. 2016. Dental topography and diets of platyrrhine primates. Historical Biology 2016;0(0):1-12. Available from, DOI: 10.1080/08912963.2016.1255737

Zliobaite, I., Rinne, J., Tóth, A., Mechenich, M, Liu, L., Behrensmeyer, A.K., and Fortelius, M. 2016. Herbivore teeth predict climatic limits in Kenyan ecosystems. PNAS 113: 12751-12756. doi: 10.1073/pnas.1609409113

Raia P, Carotenuto F, Mondanaro A, Castiglione S, Passaro F, Saggese F, Melchionna M, Serio C, Alessio L, Silvestro D & M Fortelius. 2016. Progress to extinction: increased specialisation causes the demise of animal clades. Scientific Reports 6:30965. doi: 10.1038/srep30965

Karme A, Rannikko J, Kallonen A, Clauss M, Fortelius M. 2016. Mechanical modelling of tooth wear. J. R. Soc. Interface 13: 20160399. doi: 10.1098/rsif.2016.0399

Fortelius, M., Zliobaite, I, Kaya, F., Bibi, F., Bobe, R., Leakey, L., Leakey, M.G., Patterson, D., Rannikko, J. & Werdelin, L. 2016. An ecometric analysis of the fossil mammal record of the Turkana Basin. Philosophical Transactions of the Royal Society B 371:20150232. doi:10.1098/rstb.2015.0232

Fortelius M, Geritz S, Gyllenberg M, Raia P, Toivonen J. 2015 Modeling the population-level processes of biodiversity gain and loss at geological timescales. Am. Nat. 186, 742 – 754. doi:10.1086/ 683660

Arppe, L, Kaakinen, A, Passey, B H & M Fortelius. 2015. Small mammal tooth enamel carbon isotope record of C4 grasses in late Neogene China. Global and Planetary Change 133. DOI: 10.1016/j.gloplacha.2015.09.003

Raia, P., Passaro, F., Carotenuto, F., Maiorino, L., Piras, P., Teresi, L., Meirin, S., Itescu, Y., Novosolov, M., Baiano, R.M., Fortelius, M. 2015. Cope’s Rule and the universal scaling law of ornament complexity. The American Naturalist 186, electronic publication ahead of print.

Fortelius M., Geritz S., Gyllenberg M., Toivonen, J. 2015. Adaptive dynamics on an environmental gradient that changes over a geological time-scale. Journal of Theoretical Biology 376: 91-104.

Sukselainen, L., Fortelius, M., Harrison, T. 2015. Co-occurrence of pliopithecoid and hominoid primates in the fossil record: An ecometric analysis. Journal of Human Evolution xxx:1-17. Online publication: 14-MAY-2015, DOI: 10.1016/j.jhevol.2015.04.009

Kaakinen A., Abdul Aziz, H., Passey B.H., Zhang Z., Liu L., Salminen J., Wang, L., Krijgsman W., Fortelius M. 2015. Age and stratigraphic context of Pliopithecus and associated fauna from Miocene sedimentary strata at Damiao, Inner Mongolia, China. Journal of Asian Earth Sciences 100 (2015) 78–90.

Saarinen, J.J., Boyer, A.G., Brown, J.H, Costa, D.P., Ernest, S.K.M., Evans, A.R., Fortelius, M., Gittleman, J.L., Hamilton, M.J., Harding, L.E., Lintulaakso, K., Lyons, S.K., Okie, J.G., Sibly, R.M., Stephens, P.R., Theodor, J., Uhen, M.D., Smith, F.A. 2014. Patterns of maximum body size evolution in Cenozoic land mammals: eco-evolutionary processes and abiotic forcing. Proceedings of the Royal Society B 281, doi: 10.1098/rspb.2013.2049.

Müller J., Clauss M., Codron D., Schulz E., Hummel J., Fortelius M., Kircher P., Hatt J.-M. 2014. Growth and wear of incisor and cheek teeth in domestic rabbits (Oryctolagus cuniculus) fed diets of different abrasiveness. Journal of Experimental Zoology A 321: 283-298.

Barnosky, A.D., Hadly, E. A., Dirzo, R., Fortelius, M. & Stenseth, N.C. 2014.  Translating science for decision makers to help navigate the Anthropocene.  The Anthropocene Review 1:160–170. doi: 10.1177/2053019614533670

Fortelius, M., Eronen, J.T., Kaya, F., Tang, H., Raia, P. & Puolamäki, K. 2014. Evolution of Neogene mammals in Eurasia: environmental forcing and biotic interactions. Annual Reviews of Earth and Planetary Science 42:579–604, doi: 10.1146/annurev-earth-050212-124030.

Barnosky A.D., Brown J.H., Daily G.C., et al. 2014. Introducing the Scientific Consensus on Maintaining Humanity's Life Support Systems in the 21st Century: Information for Policy Makers. The Anthropocene Review 1:78-109.

Cardoso, P.M., Rigal, F., Carvalho, J.C., Fortelius, M., Borges, P.A.V., Podani, J., Schmera, D. 2014. Partitioning taxon, phylogenetic and functional beta diversity into replacement and richness difference components. Journal of Biogeography 41:749–761.

Wang, X., L.F.  Flynn and M. Fortelius (Eds). 2013. Fossil Mammals of Asia. Neogene terrestrial biostratigraphy and chronology. Columbia University Press. 732 pp.

Raia, P. & Fortelius, M. 2013.  Cope’s Law of the Unspecialized, Cope’s Rule, and weak directionality in evolution. Evolutionary Ecology Research 15: 747–756. 

Nummela S., Pihlström, H., Puolamäki, K., Fortelius, M., Hemilä, S., Reuter, T. 2013. Exploring the mammalian sensory space: co-operations and trade-offs among senses. Journal of Comparative Physiology. A, Neuroethology, Sensory, Neural, and Behavioral Physiology 199:1077-1092.

Okie J.G., Boyer A.G., Brown J.H., Costa D.P., Ernest S.K.M., Evans A.R., Fortelius M., Gittleman J.L., Hamilton M.J., Harding L.E., Lintulaakso K., Lyons S.K., Saarinen J.J., Smith F.A., Stephens P.R., Theodor J., Uhen M.D., Sibly R.M. 2013. Effects of allometry, productivity and lifestyle on rates and limits of body size evolution. Proceedings of the Royal Society B 280: 20131007 (online June 13).

Zhang, Z., Kaakinen, A. P., Liu, L., Lunkka, J.P., Sen, S., Gose, W.A., Qiu, Z. Zheng, S.  & Fortelius, M. 2013. Mammalian biochronology of the Late Miocene Bahe Formation. Pp. 187-202 in Wang, X., Flynn, L.J.  & Fortelius, M (Eds), Fossil Mammals of Asia, New York: Columbia University Press.

Kaakinen, A. P., Passey, B. H., Zhang, Z., Pesonen, L., Liu, L. & Fortelius, M. 2013 Stratigraphy and paleoecology of the classical dragon bone localities of Baode County, Shanxi Province, China. Pp. 203-217 in Wang, X., Flynn, L.J.  & Fortelius, M (Eds), Fossil Mammals of Asia, New York: Columbia University Press.

Mirzaie Ataabadi, M., Liu, L., Eronen, J. T., Bernor, R. & Fortelius, M. 2013. Continental Scale Patterns in Neogene Mammal Community Evolution and Biogeography: A Europe-Asia Perspective. Pp. 629-655 in Wang, X., Flynn, L.J.  & Fortelius, M (Eds), Fossil Mammals of Asia, New York: Columbia University Press.

Hadly E.A., Barnosky A.D., Fortelius M., Stenseth N.C. 2013. Getting the word out on biosphere crisis. Nature 497:565. 

Fortelius, M. 2013. Commentary: The grassiness of all flesh. Journal of Biogeography 40: 1213–1214.

Uhen, M. D., Barnosky, A. D., Bills, B., Blois, J., Carrano, M. T., Carrasco, M. A., Erickson, G. M., Eronen, J. T., Fortelius, M., Graham, R. W., Grimm, E. C., O'Leary, M. A., Mast, A., Piel, W. H., Polly, P. D. & Säilä, L. K. 2013. From card catalogs to computers: Databases in vertebrate paleontology. Journal of Vertebrate Paleontology. 33:13-28.

Raia P, Carotenuto F, Passaro F, Piras P, Fulgione D, Werdelin L, Saarinen J, Fortelius M. 2013. Rapid action in the Palaeogene: the relationship between phenotypic and taxonomic diversification in Coenozoic mammals. Proceedings of the Royal Society B 280: 2012-2244.

Kaiser, T.M., Müller, D.W.H., Fortelius, M., Schultz, E., Codron, D. and Clauss, M. 2013. Hypsodonty and tooth facet development in relation to diet and habitat in herbivorous ungulates: implications for understanding tooth wear. Mammal Review 43:34–46.

Tang, H., Eronen, J. T., Micheels , A., Ahrens, B. & Fortelius, M. 2012. Asynchronous responses of East Asian and Indian summer monsoons to mountain uplift shown by regional climate modelling experiments. Climate Dynamics 40:1531-1549.

Eronen, J.T., Fortelius, M., Micheels, A., Portmann, F.T., Puolamäki, K. & Janis, C.M. 2012. Neogene aridification of the Northern Hemisphere. Geology 40:823-826.

Liu, L., Puolamäki, K., Eronen, J. T., Mirzaie Ataabadi, M., Hernesniemi, E. & Fortelius, M. 2012 Dental functional traits of mammals resolve productivity in terrestrial ecosystems past and present. Proceedings of the Royal Society B 279:2793-2799.

Wilson, G. P., Evans, A. R., Corfe, I. J., Smits, P. D., Fortelius, M. & Jernvall, J. 2012. Adaptive radiation of multituberculate mammals before the extinction of dinosaurs. Nature 483:457-460.

Heikinheimo, H., Eronen, J. T., Sennikov, A., Preston, C., Oikarinen, E., Uotila, P., Mannila, H. & Fortelius, M. 2012. Convergence in the distribution patterns of Europe's plants and mammals is due to environmental forcing. Journal of Biogeography 39:1633-1644.

Barnosky, A.D., Hadly, E.A., Bascompte, J., Berlow, E. L., Brown, J.H., Fortelius, M., Getz, W.M., Harte, J., Hastings, A., Marquet, P.A., Martinez, N.D., Mooers, A., Roopnarine, P., Vermeij, G., Williams, J.W., Gillespie, R., Kitzes, J., Marshall, C., Matzke, N., Mindell, D.P., Revilla, E. & Smith, A.B. 2012. Approaching a state shift in Earth’s biosphere. Nature 486:52-58.

Evans, A.R., Jones, D., Boyer, A.G., Brown, J.H., Costa, D.P., Dayan, T., Ernest, S.K.M.,  Fitzgerald, E.M.G. Fortelius, M.  Gittleman, J.L. Hamilton, M.J., Harding, L.E.Lintulaakso, K., Lyons, S.K., Okie, J.G. Saarinen, J.J., Sibly, R.M.,  Smith, F.A., Stephens, P.R., Theodor, J., and Uhen, M.D. 2012. Maximum rates of body size macroevolution in mammals. PNAS 109:4187-4190.

Raia P., Carotenuto F., Passaro F., Fulgione D. and Fortelius, M. 2012. Ecological specialization in fossil mammals explains Cope's rule. American Naturalist 179:328-337.

Eronen, J.T., Puolamäki, K., Heikinheimo, H., Lokki, H., Venäläinen, A., Mannila, H. and Fortelius, M. 2011. The effect of scale, climate and environment on species richness and spatial distribution of Finnish birds. Annales Zoologici Fennici 48:257-274.

Eronen, J. T., Evans, A. R., Fortelius, M. & Jernvall, J. 2011. Genera are often better than species for detecting evolutionary change in the fossil record: A reply to Salesa et al. Evolution 65:1514-1516.

Zhang, Z.Q., Wang, L.H., Kaakinen, A., Liu, L.P. and Fortelius, M. 2011. Miocene mammalian faunal succession from Damiao, central Nei Mongol and the environmental changes. Quaternary Sciences 31:608-613.

Deng, T., X. Wang, M. Fortelius, Q. Li, Y. Wang, Z.J. Tseng, G.T. Takeuchi, J. E. Saylor, L.K. Säilä, G. Xie. 2011. Out of Tibet: Pliocene woolly rhino suggests high-plateau origin of Ice Age megaherbivores. Science 333:1285-1288.

Flynn, L.F., X. Wang and M. Fortelius. 2011. Neogene terrestrial mammalian biostratigraphy and chronology in Asia: a special issue [Preface]. Vertebrata PalAsiatica 49 (3):253-256.

Wang, X. G. Xie, Z. Qiu,  Q. Li Z.J. Tseng, G.T. Takeuchi B. Wang,  M. Fortelius, A. Rosenström-Fortelius, H.  Wahlquist. Early explorations of Qaidam Basin (Tibetan Plateau) by Birger Bohlin - reconciling classic vertebrate fossil localities with modern biostratigraphy. Vertebrata PalAsiatica 49:285-310.

Tang, H., Eronen, J. T., Micheels, A. & Fortelius, M. 2011.  A regional climate model experiment to investigate the Asian monsoon in the Late Miocene. Climate of the Past 7:847-868.

Micheels, A., Bruch, A., Eronen, J. T., Fortelius, M., Harzhauser, M., Utescher, T. & Mosbrugger, V. 2011. Analysis of heat transport mechanisms from a Late Miocene model experiment with a fully-coupled atmosphere–ocean general circulation model. Palaeogeography, Palaeoclimatology, Palaeoecology 304:337-350.

Raia, P., F. Carotenuto, J. T. Eronen and M. Fortelius. 2011. Longer in the tooth, shorter in the record? The evolutionary correlates of hypsodonty in Neogene ruminants. Proceedings of the Royal Society B 278, p. 3474-3481.

Kallio, A., Puolamäki, K., Fortelius, M., and Mannila, H., 2011. Correlations and co-occurrences of taxa: the role of temporal, geographic, and taxonomic restrictions. Palaeontologia Electronica Vol. 14, Issue 1; 4A:14p;

Hernesniemi, E., I.X. Giaourtsakis, A. Evans and M. Fortelius. 2011. Rhinocerotidae. Pp. 275-294 in: T. Harrison (Ed), Paleontology and geology of Laetoli: human evolution in context. Vol. 2: Fossil hominins and the associated fauna. Springer.

Hernesniemi, E., Blomstedt, K. & Fortelius, M. 2011. Multi-view stereo 3D reconstruction of lower molars of Recent and Pleistocene rhinoceroses for mesowear analysis. Palaeontologia Electronica 14, 2, 15 pp.

Wang, X., Xie, Z., Qiu, Z., Li, Q., Tseng, Z. J., Takeuchi, G. T., Wang, B., Fortelius, M., Rosenström-Fortelius, A. & Wahlquist, H. 2011. Early explorations of Qaidam Basin (Tibetan Plateau) by Birger Bohlin - reconciling classic vertebrate fossil localities with modern biostratigraphy. Vertebrata PalAsiatica 49:285-310.

Mirzaie Ataabadi, M., Mohammadalizadeh, J., Zhang, Z., Watabe, M., Kaakinen, A. P. & Fortelius, M. 2011. Late Miocene large mammals from Ivand (Northwestern Iran). Geodiversitas. 33:709-728.

Polly, P.D., J.T. Eronen, M. Fred, G.P. Dietl, V. Mosbrugger, C. Scheidegger, D.C. Frank, J. Damuth, N.C. Stenseth, and M. Fortelius.  2011. History Matters: ecometrics and integrative climate change biology.  Proceedings of the Royal Society B, 278: 1131-1140.

Saarinen, J., E. Oikarinen, M. Fortelius, and H. Mannila. 2010. The living and the fossilised  –  how well do unevenly distributed points capture the faunal information in a grid? Evolutionary Ecology Research 12: 363–376.

Eronen, J.T., Puolamäki, K., Liu, L., Lintulaakso, K., Damuth, J., Janis, C., and Fortelius, M. 2010. Precipitation and large herbivorous mammals, part I: Estimates from present-day communities. Evolutionary Ecology Research 12, 217-233.

Eronen, J.T., Puolamäki, K., Liu, L., Lintulaakso, K., Damuth, J., Janis, C., and Fortelius, M. 2010. Precipitation and large herbivorous mammals , part II: Application to fossil data. Evolutionary Ecology Research 12, 235-248.

Eronen, J.T. , Evans, A.R., Jernvall, J., Fortelius, M. 2010. The impact of regional climate on the evolution of mammals. A case study using fossil horses. Evolution 64, 398-408. doi:10.1111/j.1558-5646.2009.00830.x

Eronen, J.T., Polly, P.D., Fred, M., Damuth, J., Frank, D.C., Mosbrugger, V., Scheidegger, C., Stenseth, N.C., Fortelius, M. 2010. Ecometrics: The traits that bind the past and present together. Integrative Zoology 5: 88-101.

Smith, F. A., Boyer, A. G., Brown, J. H., Costa, D. P., Dayan, T., Ernest, S. K. M., Evans, A. R., Fortelius, M., Gittleman, J. L., Hamilton, M. J., Harding, L. E.,Lintulaakso, K., Lyons, S. K., McCain, C., Okie, J. G., Saarinen, J. J., Sibly, R. M., Stephens, P. R., Theodor, J. & Uhen, M. D. 2010. The evolution of maximum body size of terrestrial mammals. Science. 330:1216-1219

Eronen, J. T.,  M. Mirzaie Ataabadi, A. Micheels, A. Karme, R. L. Bernor, and M. Fortelius. 2009. Distribution history and climatic controls of the Late Miocene Pikermian chronofauna. PNAS 106 (29):11867–11871.

Liu, L., J. T. Eronen, M. Fortelius. 2009.  Significant mid-latitude aridity in the middle Miocene of East Asia. Palaeogeography, Palaeoclimatology, Palaeoecology 279:201-206.

Liow, L. H., M. Fortelius; K. Lintulaakso; H. Mannila & N. C. Stenseth. 2009. Lower extinction risk in sleep-or-hide mammals. The American Naturalist 178:264-272.