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A complete list of publications by Scott Steppan can be found on his CV.
Please note: The copyright of these articles is with their respective publishers. By downloading an article, you agree to limit the use of the pdf file to printing of single copies for personal research and study. You may not modify the files in any way, nor use them for commercial purposes.


2022 Bangs, M. R. and Steppan, S. J. A rodent anchored hybrid enrichment probe set for a range of phylogenetic utility: From order to species. Molecular ecology resources 22 (4), 1521-1528DOIPDF
2022 Teta, P., Jayat, J. P., Steppan, S. J., Ojeda, A. A., Ortiz, P. E., Novillo, A., Lanzone, C., and Ojeda, R. A. Uncovering cryptic diversity does not end: a new species of leaf-eared mouse, genus Phyllotis (Rodentia, Cricetidae), from Central Sierras of Argentina. MammaliaDOIPDF
2022 Saltzberg, C. J., Walker, L. I., Chipps-Walton, L. E., Costa, B., Spotorno, A. E., and Steppan, S. J. Comparative Quantitative Genetics of the Pelvis in Four-Species of Rodents and the Conservation of Genetic Covariance and Correlation Structure. Evolutionary Biology 49 (1): 71-83DOI
2022 Steppan, S. J., Meyer, A. A., Barrow, L. N., Alhajeri, B. H., Al-Zaidan, A. S. Y., Gignac, P. M., and Erickson, G. M. Phylogenetics And The Evolution Of Terrestriality In Mudskippers (Gobiidae: Oxudercinae). Molecular Phylogenetics and Evolution 169: 107416DOIPDF
2022 Steppan, S. J., Bowen, T., Bangs, M. R., Farson, M., Storz, J. F., Quiroga-Carmona, M., D’Elía, G., Vimercati, L., Ortiz, C. D., Zimmerman, G., and Schmidt, S. K.Evidence of a population of leaf-eared mice Phyllotis vaccarum above 6,000 m in the Andes and a survey of high-elevation mammals. Journal of Mammology DOIPDF


2021 Jayat, J. P., Teta, P., Ojeda, A. A., Steppan, S. J., Osland, J. M., Ortiz, P. E., Novillo, A., Lanzone, C., Ojeda, R. A.The Phyllotis xanthopygus complex (Rodentia, Cricetidae) in central Andes, systematics and description of a new species. Zoologica Scripta. In press.
2021 Percequillo, R. A., do Prado, J. R., Abreu, E. F., Dalapicolla, J., Pavan, A. C., Chiquito, E. A., Brennand, P., Steppan, S. J., Lemmon, A. R., Lemmon, E. M., Wilkinson, M. Tempo and mode of evolution of oryzomyine rodents (Rodentia, Cricetidae, Sigmodontinae): A phylogenomic approach. Molecular Phylogenetics and Evolution. 159DOIPDF


2020 Storz, J. F., Quiroga-Carmona, M., Opazo, J. C., Bowen, T., Farsone, M., Steppan, S. J., and D’Elía, G. Discovery of the world’s highest-dwelling mammal. Proceedings of the National Academy of Science. DOIPDF


2019 Rowe, K. C., Achmadi, A. S., Fabre, P-H., Schenk, J. J., Steppan, S. J., and Esselstyn, J. A. Oceanic islands of Wallacea as a source for dispersal and diversification of murine rodents. Journal of Biogeography. 46:2752–2768. DOIPDF


2018 Young, G. R., Yap, M. W., Michaux, J. R., Steppan,, S. J., and Stoye, J. P. Evolutionary journey of the retroviral restriction gene Fv1. Proceedings of the National Academy of Science. DOIPDF
2018 Alhajeri, B. H. and Steppan, S. J. Interspecific variation in skull morphology of rodents along aridity clines: a phylogenetic test of desert adaptation. Journal of Mammalogy. 99 (5):117–1216. 
2018 Alhajeri, B. H. and Steppan, S. J. Community structur e in ecological assemblages of desert rodents. Biological Journal of the Linnean Society. 124, 308–318.
2018 Schenk, J. J., and Steppan, S. J. The role of geography in adaptive radiation. American Naturalist. 192 (4): 415-431.
2018 Alhajeri, B.H. and Steppan, S.J. Disparity and Evolutionary Rate Do Not Explain Diversity Patterns in Muroid Rodents (Rodentia: Muroidea). Evolutionary Biology.DOI
2018 Alhajeri, B.H. and Steppan, S.J. Ecological and ecomorphological specialization are not associated with diversification rates in muroid rodents (Rodentia: Muroidea). Evolutionary Biology.DOIPDF
2018 Heaney, L. R., Kyriazis, C. C., Balete, D. S., Steppan, S. J., and Rickart, E. A. How small an island? Speciation by endemic mammals (Apomys, Muridae) on an oceanic Philippine island. Journal of Biogeography.DOIPDF
2018 Heaney, L. R., Kyriazis, C. C., Balete, D. S., Steppan, S. J., and Rickart, E. A. How small an island? Speciation by endemic mammals (Apomys, Muridae) on an oceanic Philippine island. Journal of Biogeography.DOIPDF


2017 Steppan, S. J. and Schenk, J. J. Muroid rodent phylogenetics: 900-species tree reveals increasing diversification rates. PLoSOne. 12(8): e0183070.DOIPDF
2017 Wray, K. P., Means, D. B., and Steppan, S. J. Revision of the Eurycea quadridigitata (Holbrook 1842) complex of dwarf salamanders (Caudata: Plethodontidae: Hemidactyliinae) with a description of two new species. Herpetological Monographs. 31 (1): 18-46. DOIPDF
2017 Wray, K. P., and Steppan, S. J. Historical biogeography and diversification in a major lineage of salamanders (Plethodontidae: Spelerpinae) Journal of Biogeography. 44:797-809. DOIPDF


2016 Heaney, L.R., Balete, D.S., Duya, M.R.M., Duya, M.V., Jansa, S. A., Steppan, S.J., Rickart, E.A. Doubling diversity: a cautionary tale of previously unsuspected mammalian diversity on a tropical oceanic island. Frontiers in Biogeography: 1-19.DOIPDF
2016 Martin, S.A., Alhajeri, B. H., and Steppan, S.J. Dietary adaptations in teeth of murine rodents (Muridae). Biological Journal of the Linnean Society. 119: 766-784.DOIPDF
2016 Alhajeri, B. H. and Steppan, S. J. Association between climate and body size in rodents (Order: Rodentia): A phylogenetic test of Bergmann’s rule. Mammalian Biology. 81:219-26.DOIPDF
2016 Gingerich, T.J., Stumpo, D.J., Lai, W.S., Randall, T.A., Steppan, S.J., and Blackshear, P.J. Rapid Emergence and Evolution of Zfp36l3. Molecular Phylogenetics and Evolution. 94: 518-530.DOIPDF
2016 Alhajeri, B. H., Schenk, J. J., and Steppan, S. J. Ecomorphological diversification following continental colonization in muroid rodents: a phenotypic test of the ecological opportunity model of adaptive radiation. Biological Journal of the Linnean Society. 117: 463–481.DOIPDF


2015 Herrera, N. D., ter Poorten, J. J., Bieler, R., Mikkelsen, P. M., Strong, R. E., Jablonski, D., and Steppan, S. J. Molecular phylogenetics and historical biogeography amid shifting continents in the cockles and giant clams (Bivalvia: Cardiidae). Molecular Phylogenetics and Evolution. 93:94-106.DOIPDF
2015 Alhajeri, B. H., Hunt, O. J., and Steppan, S. J. Molecular systematics of gerbils and deomyines (Rodentia: Gerbillinae, Deomyinae) and a test of desert adaptation in the tympanic bulla. Journal of Zoological Systematics and Evolutionary Research. 53(4): 312-330. DOIPDF
2015 Steppan, S. J. and Ramirez-Baca, O. Genus Phyllotis. In Mammals of South America. (Patton, J. L., Pardiñas, U.F.J., & D’Elia, G. ed.). University of Chicago Press. Pp 535-555.Link
2015 Polly, P. D., Cardini, A., Davis, E.B., and Steppan, S. J. Marmot evolution and global change in the last 10 million years. In Evolution of the Rodents: Advances in Phylogeny, Palaeontology and Functional Morphology, (Cox, P. and Hautier, L. eds.) Cambridge Univ. Press. Pp 246-276. DOIPDF
2015 Justiniano, R., Schenk, J.J., Balete, D.S., Rickart, E.A., Esselstyn, J.A. Heaney, L.R., and Steppan, S. J. Testing diversification models of endemic Philippine forest mice (Apomys) with nuclear phylogenies across elevational gradients reveals repeated colonization of isolated mountain ranges. Journal of Biogeography. 42(1):51-64.DOIPDF

2014 - 2010

2014 Schenk, J.J., and Steppan, S. J. Too long to read: Assessing the motivation behind graduate student attendance in reading groups. Journal of College Science Teaching. 44(2):64-69. DOIPDF
2014 Heaney, L.R., Balete, D.S., Veluz, M.J., Steppan, S. J., Esselstyn, J.A., Pfeiffer, A.W., and Rickart, E.A. Two new species of Philippine forest mice (Apomys, Muridae, Rodentia) from Lubang and Luzon Islands, with a redescription of Apomys sacobianus Johnson, 1962. Proceedings of the Biological Society of Washington. 126(4):395–413 DOIPDF
2013 Schenk, J.J., Rowe, K.C., and Steppan, S. J. Ecological opportunity and incumbency in the diversification of repeated continental colonizations by muroid rodents. Systematic Biology. 62(6):837–864. DOIPDF
2012 Ortiz, P. E., Jayat, J. P., and Steppan, S. J. A new fossil Phyllotini (Rodentia: Sigmodontinae) from Late Pliocene in the Andes of northwestern Argentina. Journal of Vertebrate Paleontology 32 (6): 1429-1441. DOIPDF
2012 Erickson, G.M., Gignac, P. M., Steppan, S. J., Lappin, A. K., Vliet, K. A., Brueggen, J. D., Inouye, B. D., Kledzik, D., Webb, G. J. W. Insights into the ecology and evolutionary success of crocodilians revealed through bite force and tooth pressure experimentation. PLoS ONE 7(3): e31781. DOIPDF
2011 Steppan, S. J., Kenagy, G. J., Zawadzki, C., Robles, R., Lyupanova, E.A., and Hoffmann, R.S. Molecular data resolve placement of the Olympic marmot and estimate dates of trans-Beringian interchange. Journal of Mammalogy. 92(5):1028-1037. DOIPDF
2011 Burns, J. H., Faden, R. B., and Steppan, S. J. Phylogenetic studies in the Commelinaceae subfamily Commelinoideae inferred from nuclear ribosomal and chloroplast DNA sequences. Systematic Botany. 36(2):268-276. DOIPDF
2011 Lawrence R. Heaney, Danilo S. Balete, Eric A. Rickart, Phillip A. Alviola, Mariano Roy M. Duya, Melizar V. Duya, M. Josefa Veluz, Lawren VandeVrede, and Scott J. Steppan. Seven new species and a new subgenus of forest mice (Rodentia: Muridae: Apomys) from Luzon Island, Philippines. Fieldiana: Life and Earth Sciences 2: 1–60. DOIPDF
2010 Cooper, J., and Steppan, S. J. Developmental constraint on the evolution of marsupial forelimb morphology. Australian Journal of Zoology. 58: 1-15. DOIPDF
2010 van der Linde, K., Houle, D., Spicer, G., and Steppan, S. J. A supermatrix analysis of the family Drosophilidae. Genetics Research. 92: 25-38 . DOIPDF


2008 Harshman, J., Braun, E. L., Braun, M. J., Huddleston, C. J., Bowie. R. C. K., Chojnowski, J. L., Hackett, S. J., Han, K-L., Kimball, R. T, Marks, B. D., Miglia, K. J., Moore, W. S., Reddy, S., Sheldon, F. H., Steadman, D. W., Steppan, S. J., Witt, C. J., and Yuri, T. Phylogenomic evidence for multiple losses of flight in ratite birds. Proceedings of the National Academy of Science 105 (36): 13462-13467. DOIPDF
2008 Steppan, S. J. Aligning the spaces: a comment on Polly - Developmental dynamics and G-matrices. Evolutionary Biology. 35(2):108-110. DOIPDF
2008 Good, J. M., Hird, S., Demboski, J. R., Steppan, S. J., Martin-Nims, T.R., and Sullivan, J. Ancient hybridization and mitochondrial capture between two distantly related species of chipmunks (Tamias: Rodentia). Journal of Molecular Ecology. 17(5):1313-1327.DOIPDF
2008 Rowe, K., Reno, M. L., Richmond, D. M., Adkins, R. M., and Steppan, S. J. Pliocene colonization and adaptive radiations in Australia and New Guinea (Sahul): Multi-locus systematics of the Old Endemic rodents (Muroidea: Murinae). Molecular Phylogenetics and Evolution. 47: 84-101.DOIPDF
2007 Steppan, S. J., Ramirez, O., Banbury, J., Huchon, D., Pacheco, V., Walker, L, and Spotorno, A.O.. A molecular reappraisal of the systematics of the leaf-eared mice Phyllotis and their relatives. Pp. 799-820 in The quintessential naturalist: honoring the life and legacy of Oliver P. Pearson (Kelt, D. A., E. P. Lessa, J. A. Salazar-Bravo, and J. L. Patton, eds.). University of California Publications in Zoology 134.DOIPDF
2005 Steppan, S. J., Adkins, R. M., Spinks, P. Q., and Hale, C. Multigene phylogeny of the Old World mice Murinae reveals distinct geographic lineages and the declining utility of mitochondrial genes compared to nuclear genes. Molecular Phylogenetics and Evolution. 37:370-388.DOIPDF
2005 Steppan, S. J., Seeing the Forest for the Trees. Book review of “Assembling the Tree of Life.” Science. 307:677-678.DOIPDF
2004 Steppan, S. J., Adkins, R.M., and Anderson, J. Phylogeny and divergence date estimates of murid rodents based on multiple nuclear genes. Systematic Biology. 53(4):533-553.DOIPDF
2004 Holliday, J. A. and Steppan, S. J. Evolution of hypercarnivory: the effect of specialization on morphological and taxonomic diversity. Paleobiology. 30(1):108-128.DOIPDF
2004 Steppan, S. J. Phylogenetic comparative analysis of multivariate data. In Phenotypic Integration. (eds., Piggliuci, M and Preston, K. A.). Oxford Univ. Press.LINKPDF
2004 Steppan, S. J., Storz, B. L., and Hoffmann, R. S. Nuclear DNA phylogeny of the squirrels (Mammalia: Rodentia) and the evolution of arboreality from c-myc and RAG1. Molecular Phylogenetics and Evolution. 30:703-719.DOIPDF
2003 Steppan, S. J., Zawadzki, C., and Heaney, L. R. Molecular phylogeny of the endemic Philippine rodent Apomys (Muridae) and the dynamics of diversification in an oceanic archipelago. Biological Journal of the Linnean Society. 80:699-715 .DOIPDF
2002 Steppan, S. J., Houle, D., and Phillips, P.C. Comparative quantitative genetics: evolution of the G matrix. Trends in Ecology and Evolution. 17:320-327.DOIPDF
2002 Kuch, M., Rohland, N., Betancourt, J., Lattore, C., Steppan, S. J., and Poinar, H.N. Molecular Analysis of a 11,700-yr old rodent midden from the Atacama Desert Chile. Molecular Ecology. 11:913-924.DOIPDF
2000 Steppan, S. J. Flexural stiffness patterns of butterfly wings. Journal of Research on the Lepidoptera. 35:61-77.PDF
2000 Steppan, S. J. and Sullivan, J. The emerging statistical perspective in systematics and the status of Andalgalomys (Rodentia: Sigmodontinae): a comment on Mares and Braun. Journal of Mammalogy. 81(1):260-270.DOIPDF
2000 Ortiz, P.E., Pardiñas, U.F.J. and Steppan, S. J. A new extinct phyllotine (Rodentia: Muridae) from the Pleistocene of Argentina and a consideration of the phylogenetic relationships of the “Reithrodon group.” Journal of Mammalogy. 81(1):37-51.DOIPDF
1999 Steppan, S. J., Akhverdyan, M. R., Lyapunova, E.A., Fraser, D.G., Vorontsov, N.N., Hoffmann, R.S., and Braun, M.J. Molecular phylogeny of the marmots (Rodentia: Sciuridae): tests of evolutionary and biogeographic hypotheses. Systematic Biology.48(4):715-734.DOIPDF
1998 Steppan, S. J. and U. Pardiñas. Two new fossil murids from the Pleistocene of Argentina: phylogeny and paleoecology. Journal of Vertebrate Paleontology. 18(3):640-649.DOIPDF
1998 Steppan, S. J. Phylogenetic relationships and species limits within Phyllotis (Rodentia: Sigmodontinae): concordance between mtDNA sequence and morphology. Journal of Mammalogy. 79(2):573-593.DOIPDF
1997 Steppan, S. J. Phylogenetic analysis of phenotypic covariance structure. II. Reconstructing matrix evolution. Evolution. 51(2): 587-594.DOIPDF
1997 Steppan, S. J. Phylogenetic analysis of phenotypic covariance structure. I. Contrasting results from matrix correlation and Common Principal Component analyses. Evolution. 51(2): 571-586. DOIPDF
1996 Steppan, S. J. A new species of Holochilus (Rodentia: Sigmodontinae) from the middle Pleistocene of Bolivia and its phylogenetic significance. Journal of Vertebrate Paleontology. 16(3):522-530.DOIPDF
1995 Steppan, S. J. Revision of the tribe Phyllotini (Rodentia: Sigmodontinae) with a phylogenetic hypothesis for the Sigmodontinae. Fieldiana: Zoology. n.s. 80:1-112.DOIPDF
1993 Steppan, S. J. Phylogenetic relationships among the Phyllotini (Rodentia: Sigmodontinae) using morphological characters. Journal of Mammalian Evolution. 1(3):187-213.DOIPDF
1991 Steppan, S. J. Geographic distribution of flower morphological traits in subspecies of Lotus scoparius. Journal of Biogeography. 18:321-331.DOIPDF

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