Molecular phylogeny of treeshrews (Mammalia: Scandentia) and the timescale of diversification in Southeast Asia Trina E. Roberts a,b,⇑ , Hayley C. Lanier a,c , Eric J. Sargis d,e , Link E. Olson a,f a University of Alaska Museum, University of Alaska Fairbanks, 907 Yukon Dr., Fairbanks, AK 99775, USA b National Evolutionary Synthesis Center, 2024 W. Main St., Suite A200, Durham, NC 27705, USA c Department of Ecology and Evolutionary Biology, Museum of Zoology, University of Michigan, 1109 Geddes Ave., Ann Arbor, MI 48109, USA d Department of Anthropology, Yale University, P.O. Box 208277, New Haven, CT 06520, USA e Divisions of Vertebrate Zoology and Vertebrate Paleontology, Yale Peabody Museum of Natural History, P.O. Box 208118, New Haven, CT 06520, USA f Institute of Arctic Biology, University of Alaska Fairbanks, 902 N. Koyukuk Dr., Fairbanks, AK 99775, USA article info Article history: Received 8 December 2010 Revised 11 April 2011 Accepted 26 April 2011 Available online 1 May 2011 Keywords: Scandentia Tupaiidae Southeast Asia Borneo Miocene Treeshrews abstract Resolving the phylogeny of treeshrews (Order Scandentia) has historically proven difficult, in large part because of access to specimens and samples from critical taxa. We used ‘‘antique’’ DNA methods with non-destructive sampling of museum specimens to complete taxon sampling for the 20 currently recog- nized treeshrew species and to estimate their phylogeny and divergence times. Most divergence among extant species is estimated to have taken place within the past 20 million years, with deeper divergences between the two families (Ptilocercidae and Tupaiidae) and between Dendrogale and all other genera within Tupaiidae. All but one of the divergences between currently recognized species had occurred by 4 Mya, suggesting that Miocene tectonics, volcanism, and geographic instability drove treeshrew diver- sification. These geologic processes may be associated with an increase in net diversification rate in the early Miocene. Most evolutionary relationships appear consistent with island-hopping or landbridge colonization between contiguous geographic areas, although there are exceptions in which extinction may play an important part. The single recent divergence is between Tupaia palawanensis and Tupaia moellendorffi, both endemic to the Philippines, and may be due to Pleistocene sea level fluctuations and post-landbridge isolation in allopatry. We provide a time-calibrated phylogenetic framework for answering evolutionary questions about treeshrews and about evolutionary patterns and processes in Euarchonta. We also propose subsuming the monotypic genus Urogale, a Philippine endemic, into Tupaia, thereby reducing the number of extant treeshrew genera from five to four. Ó 2011 Elsevier Inc. All rights reserved. 1. Introduction With modern methods of phylogenetic inference and diver- gence date estimation, biologists can form a better picture of the past than ever before. Major advances in technology have allowed the use of historical museum specimens, often collected before the advent of DNA sequencing, in genetic studies. For taxonomic groups or geographic regions for which representative samples are particularly hard to obtain, such ‘‘antique’’ (as opposed to truly ancient) DNA from historical collections can provide an incompa- rable source of evolutionary information. An understanding of evolutionary relationships within the order Scandentia (treeshrews) has lagged behind many other groups, despite the relatively small number of currently recognized species. No systematic study has ever included all recognized tree- shrew species, both because of the group’s unstable taxonomy and because of the inaccessibility of specimens and/or samples suitable for DNA extraction. This has made it impossible to get even a mod- erately complete picture of morphological or biogeographic evolu- tion in the group. In 2005, Olson et al. published a molecular phylogeny including 16 of the 20 recognized species in the order, based on the mitochondrial 12S rRNA gene. Though some relation- ships were well supported in their phylogeny, others were unclear, and some crucial questions depended on taxa for which no fresh samples were available. Roberts et al. (2009) analyzed six nuclear genes for a subset of taxa, but pointed out that missing taxa could affect hypothesized relationships. We build on these previous re- sults by adding both additional mitochondrial DNA and additional taxa, in some cases using ‘‘antique’’ DNA from museum specimens, to present the first molecular phylogeny including all currently recognized treeshrew species. A single molecular phylogeny is no substitute for a thorough revision of the order using multiple data 1055-7903/$ - see front matter Ó 2011 Elsevier Inc. All rights reserved. doi:10.1016/j.ympev.2011.04.021 ⇑ Corresponding author address: National Evolutionary Synthesis Center, Duke Box 1044003, 2024 W. Main St., Suite A200, Durham, NC 27705, USA. Fax: +1 919 668 9198. E-mail addresses: trina.roberts@nescent.org (T.E. Roberts), hclanier@umich.edu (H.C. Lanier), eric.sargis@yale.edu (E.J. Sargis), link.olson@alaska.edu (L.E. Olson). Molecular Phylogenetics and Evolution 60 (2011) 358–372 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev