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An Analysis of Pterosaurian Biogeography: Implications for the Evolutionary History and Fossil Record Quality of the First Flying Vertebrates

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An Analysis of Pterosaurian Biogeography: Implications for the Evolutionary History and Fossil Record Quality of the First Flying Vertebrates View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Natural History Museum Repository Historical Biology An International Journal of Paleobiology ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: http://www.tandfonline.com/loi/ghbi20 An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates Paul Upchurch, Brian Andres, Richard J. Butler & Paul M. Barrett To cite this article: Paul Upchurch, Brian Andres, Richard J. Butler & Paul M. Barrett (2015) An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates, Historical Biology, 27:6, 697-717, DOI: 10.1080/08912963.2014.939077 To link to this article: http://dx.doi.org/10.1080/08912963.2014.939077 © 2014 The Author(s). Published by Taylor & View supplementary material Francis. Published online: 28 Jul 2014. Submit your article to this journal Article views: 1921 View related articles View Crossmark data Citing articles: 5 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=ghbi20 Download by: [Natural History Museum] Date: 20 April 2017, At: 07:15 Historical Biology, 2015 Vol. 27, No. 6, 697–717, http://dx.doi.org/10.1080/08912963.2014.939077 An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates Paul Upchurcha*, Brian Andresb, Richard J. Butlerc and Paul M. Barrettd aDepartment of Earth Sciences, University College London, Gower Street, London WC1E 6BT, UK; bDepartment of Geology, University of South Florida, 4202 East Fowler Avenue, SCA528, Tampa, FL 33630, USA; cSchool of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK; dDepartment of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, UK (Received 24 April 2014; accepted 24 June 2014; first published online 3 September 2014) The biogeographical history of pterosaurs has received very little treatment. Here, we present the first quantitative analysis of pterosaurian biogeography based on an event-based parsimony method (Treefitter). This approach was applied to a phylogenetic tree comprising the relationships of 108 in-group pterosaurian taxa, spanning the full range of this clade’s stratigraphical and geographical extent. The results indicate that there is no support for the impact of vicariance or coherent dispersal on pterosaurian distributions. However, this group does display greatly elevated levels of sympatry. Although sampling biases and taxonomic problems might have artificially elevated the occurrence of sympatry, we argue that our results probably reflect a genuine biogeographical signal. We propose a novel model to explain pterosaurian distributions: pterosaurs underwent a series of ‘sweep-stakes’ dispersal events (across oceanic barriers in most cases), resulting in the founding of sympatric clusters of taxa. Examination of the spatiotemporal distributions of pterosaurian occurrences indicates that their fossil record is extremely patchy. Thus, while there is likely to be genuine information on pterosaurian diversity and biogeographical patterns in the current data-set, caution is required in its interpretation. Keywords: dispersal; diversity; pterosaur; sympatry; Treefitter; vicariance 1. Introduction these and other hypotheses by applying a cladistic biogeographical analysis using Treefitter 1.2b (Ronquist After their origin in the Middle or Late Triassic, pterosaurs acquired a virtually global distribution and their remains 1998; Sanmartin and Ronquist 2004), to a recent are now known from every continent, including Antarctica phylogeny for pterosaurs (Andres et al. 2014) termed (Barrett et al. 2008; see Fossilworks and The Paleobiology here the ‘reference phylogeny’ (Figures 1 and 2), in order Database). As with dinosaurs and many other clades, to determine whether there is any statistical support for pterosaurian evolution took place against a backdrop of particular distribution patterns. Such analyses also enable profound changes in palaeogeography driven by the an assessment of the relative importance of processes such fragmentation of Pangaea, major fluctuations in sea level as vicariance, dispersal, extinction and sympatric specia- and shifts in climatic zones. It is therefore surprising that tion in pterosaurian evolution. Finally, we end with a brief there has been very little detailed study of pterosaurian discussion of the quality of the pterosaurian fossil record biogeographical history (though see Unwin 1996; Wang and future requirements and prospects for further work on et al. 2005, 2007, 2012). This neglect may reflect the the biogeographical history of this clade. intense focus on the flight mechanics of these organisms, and/or the implicit assumption that the geographical distributions of flying organisms are affected more by 2. Pterosaurian distributions through space and time specific ecological requirements rather than large-scale Below, we use the atlas of pterosaurian distributions by vicariance and coherent dispersal patterns. In this paper, Barrett et al. (2008) (with revisions based on The we present the first detailed analytical study of Paleobiology Database (http://paleobiodb.org/#/), Fossil- pterosaurian biogeographical history. First, we provide works (http://fossilworks.org/) and Brian Andres, pers. an overview of the pterosaurian fossil record, summarising obs.) to generate an overview of this group’s spatiotem- where and when particular clades are represented and poral distribution (Figures 3–7, Table 1). This review adding further information based on ghost ranges. Second, provides a framework for the analyses that follow and also we briefly review the small number of previous studies that raises several issues that we believe should be addressed have proposed hypotheses to account for aspects of the by future studies. The reader should note that there are spatiotemporal distributions of pterosaurs. Third, we test some inconsistencies between the various classifications of *Corresponding author. Email: [email protected] q 2014 The Author(s). Published by Taylor & Francis. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. The moral rights of the named author(s) have been asserted. 698 P. Upchurch et al. Figure 1. The pterosaur relationships and stratigraphical/geographical ranges used in the ‘all taxa’ Treefitter data-set. This tree is based on the cladogram presented by Andres et al. (2014) and shows the more basal portion in detail (Eupterodactyloidea has been condensed to a single branch – see Figure 2). The thick branches represent known stratigraphical ranges (based on data in The Paleobiology Database); thin branches represent estimated ghost ranges and connectors used to demarcate phylogenetic relationships. Time-sliced data-sets were derived from this tree by appropriate inclusion/exclusion of taxa. Most stratigraphical stage and taxon abbreviations are listed in the legend of Table 1. Additional abbreviations: CA, Central Asia; CO, Coniacian; EA, East Asia; Eop, Eopterosauria; EU, Europe; Euc, Euctenochasmatia; KI, Kimmeridgian; NA, North America; OX, Oxfordian; RH, Rhaetian; SA (after taxon name), South America; SA (time scale), Santonian; TU, Turonian. pterosaurs applied by Barrett et al. (2008)andin of basal pterosaurs imply the existence of at least three Fossilworks and The Paleobiology Database and the lineages during the Late Triassic (Andres et al. 2010) and as reference phylogeny (Figures 1 and 2) employed here in many as seven (Kellner 2003; Wang et al. 2008); however, the Treefitter analyses. Here, we have employed the reference phylogeny used here (Figure 1) supports the pterosaurian group names and taxonomic contents that existence of only one major ghost range during this interval. are consistent with the phylogeny presented by Andres Minimally, body fossils and ghost ranges indicate that et al. (2014). members of both the Macronychoptera and Eopterosauria were present as early as the Carnian (Figure 1), although no Triassic fossils belonging to the former clade have been 2.1 Middle and Late Triassic found to date. Thus, pterosaurs almost certainly had a pre- The sister taxon to Pterosauria within Ornithodira, the Carnian origin. The first pterosaurian remains are known Dinosauromorpha, has its earliest known body fossils in from strata of probable late Carnian–early Norian age in deposits of Anisian age (Nesbitt et al. 2010, 2013), and North America and include material assigned to Eudimor- trackways suggest that this clade dates back to the early phodon (Murry 1986; Lucas and Luo 1993; Andres 2006; Olenekian (Brusatte et al. 2011)(Figure 3). This implies Figure 3, Table 1). Other pterosaurian remains have been that the pterosaurian lineage was also present in the Middle reported from Carnian and Norian sediments in this region, Triassic, although the oldest body fossils of this clade are but these specimens are indeterminate (e.g. Hunt and Lucas Carnian in age (see below). Previous phylogenetic analyses 1993) and cannot be confirmed as pterosaurs (Andres Historical Biology 699 Figure
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