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Author's personal copy Shark tales: A molecular species-level phylogeny of sharks (Selachimorpha, Chondrichthyes) Ximena V��lez-Zuazo a,b,���, Ingi Agnarsson a a Department of Biology, University of Puerto Rico ��� Rio Piedras (UPRRP), San Juan, PR 00931-3360, PR, USA b ecOceanica, Copernico 179, Lima 41, Peru a r t i c l e i n f o Article history: Received 19 May 2010 Revised 18 November 2010 Accepted 22 November 2010 Available online 30 November 2010 Keywords: Elasmobranchii Galeomorphii Squalimorphii Taxonomy Taxon sampling a b s t r a c t Sharks are a diverse and ecologically important group, including some of the ocean���s largest predatory animals. Sharks are also commercially important, with many species suffering overexploitation and fac- ing extinction. However, despite a long evolutionary history, commercial, and conservation importance, phylogenetic relationships within the sharks are poorly understood. To date, most studies have either focused on smaller clades within sharks, or sampled taxa sparsely across the group. A more detailed spe- cies-level phylogeny will offer further insights into shark taxonomy, provide a tool for comparative anal- yses, as well as facilitating phylogenetic estimates of conservation priorities. We used four mitochondrial and one nuclear gene to investigate the phylogenetic relationships of 229 species (all eight Orders and 31 families) of sharks, more than quadrupling the number of taxon sampled in any prior study. The resulting Bayesian phylogenetic hypothesis agrees with prior studies on the major relationships of the sharks phy- logeny however, on those relationships that have proven more controversial, it differs in several aspects from the most recent molecular studies. The phylogeny supports the division of sharks into two major groups, the Galeomorphii and Squalimorphii, rejecting the hypnosqualean hypothesis that places batoids within sharks. Within the squalimorphs the orders Hexanchiformes, Squatiniformes, Squaliformes, and Pristiophoriformes are broadly monophyletic, with minor exceptions apparently due to missing data. Similarly, within Galeomorphs, the orders Heterodontiformes, Lamniformes, Carcharhiniformes, and Orectolobiformes are broadly monophyletic, with a couple of species ���misplaced���. In contrast, many of the currently recognized shark families are not monophyletic according to our results. Our phylogeny offers some of the first clarification of the relationships among families of the order Squaliformes, a group that has thus far received relatively little phylogenetic attention. Our results suggest that the genus Ech- inorhinus is not a squaliform, but rather related to the saw sharks, a hypothesis that might be supported by both groups sharing ���spiny��� snouts. In sum, our results offer the most detailed species-level phylogeny of sharks to date and a tool for comparative analyses. �� 2010 Elsevier Inc. All rights reserved. 1. Introduction In the world���s oceans, sharks are the most diverse group of large predatory animals (Fig. 1) and they play an important ecological role as primary predators of many species (Myers and Worm, 2003 Myers et al., 2007 Schindler et al., 2008 Heithaus et al., 2008 Vaudo and Heithaus, 2009) and structuring marine ecosys- tems and food webs (Libralato et al., 2005). Sharks are also commer- cially important, sought after for their meat, in particular their fins. As a result of unsustainable fishing activities and non-specific fishing gear, worldwide sharks are being decimated and many spe- cies are facing extinction (Robbins et al., 2006 Dulvy et al., 2008 Ferretti et al., 2008). Nearly 93% (468 of 504) of shark species are currently listed by the IUCN Red List (2009, www.iucnredlist.org). This list will very likely set the tone for upcoming conservation efforts and funding allocation priorities in the next years. The deci- sion on what species to conserve is mostly based on an exhaustive assessment of anthropogenic threats and resulting estimated pop- ulation decline (http://www.iucn.org/redlist, Vi�� et al., 2008). With the advent of molecular data, information of population genetic structure and connectivity has started to be considered (e.g. Mortimer and Donnelly, 2007) and recently species phylogenetic position and evolutionary distinctiveness have started to play a role in establishing conservation priorities (Faith, 1992, 2008 Steel et al., 2007 Isaac et al., 2007 Mooers et al., 2008). Information about species evolutionary history, and the status of their close relatives, can impact conservation planning, especially for those already identified as under some level of threat. 1055-7903/$ - see front matter �� 2010 Elsevier Inc. All rights reserved. doi:10.1016/j.ympev.2010.11.018 ��� Corresponding author at: Department of Biology, University of Puerto Rico ��� Rio Piedras (UPRRP), San Juan, PR 00931-3360, PR, USA. E-mail address: xvelezuazo@gmail.com (X. V��lez-Zuazo). Molecular Phylogenetics and Evolution 58 (2011) 207���217 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev