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Please Scroll Down for Article PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Journal of Systematic Palaeontology,Vol.8,Issue4,December2010,607–625 Morphological diversity and biogeography of procolophonids (Amniota: Parareptilia) a b Juan Carlos Cisneros and Marcello Ruta ∗ aDepartamento de Paleontologia e Estratigrafia, Universidade Federal do Rio Grande do Sul, Av. Bento Gonc¸alves 9500, Porto Alegre, CP 15001, 91540-000, Brasil; bDepartment of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol BS8 1RJ, UK (Received 4 June 2009; accepted 6 May 2010) A recent phylogenetic analysis of procolophonid parareptiles is used as the basis for a study of morphological diversity (disparity) in these amniotes. Disparity values are compared in three groups of procolophonids (a paraphyletic series of basal taxa and two monophyletic sister groups: procolophonines and leptopleuronines), two ecophenotypic assemblages (one based upon inferred diet – non high-fibre versus high-fibre species; the other based upon cranial sculpture – non horned versus horned species), and two temporal assemblages (Lower Triassic versus Middle and Upper Triassic). The mean disparity values are comparable in the case of temporal and ecophenotypic assemblages. High-fibre species are marginally less disparate than non high-fibre species. The combined Middle and Upper Triassic species are slightly less disparate than Lower Triassic species. Finally, horned species are only slightly more disparate than non-horned species. The paraphyletic series of basal taxa and the leptopleuronines show similar disparity values, marginally higher than those for procolophonines. Phylogenetic analysis is also used to reconstruct the biogeographical history of procolophonids. Both ancestral area analysis and dispersal–vicariance analysis show that South Africa was the most likely ancestral area for procolophonids as a whole. North China – either as a single area or in combination with Russia or South Africa – was the most likely ancestral area for the leptopleuronine–procolophonine clade. Keywords:Procolophonidae;morphospace;disparity;dispersal;ancestralarea;phylogeny Introduction 2004; Jalil & Janvier 2005; Modesto 2006; Tsuji 2006; Modesto & Damiani 2007; Muller¨ & Tsuji 2007; Reisz et al. The clade Parareptilia (see Tsuji & Muller¨ 2009 for a 2007; Cisneros 2008a, b; Muller¨ et al.2008;Sail¨ a2008,¨ novel recent definition and a synoptic overview) consists 2009a; Sues & Reisz 2008; Tsuji & Muller¨ 2008, 2009). of a diverse array of Permian and Triassic reptiles (sensu Procolophonoidea – one of the most species-rich clades Modesto & Anderson 2004) of global distribution and great of parareptiles – includes superficially lizard-like reptiles zoological significance, not least because several of them from the Permian and Triassic, diagnosed by (among other Downloaded By: [Cisneros, Juan Carlos] At: 10:54 6 November 2010 have been proposed, not without controversy, as the ances- characters) the absence of a tabular, a contribution of the tors or near-ancestors of turtles (e.g. Reisz & Laurin 1991; jugal to the ventral embayment of the posterior cheek, Laurin & Reisz 1995; Lee 1995, 1997a; see also comments an emarginated ventral margin of the temporal region in Rieppel & Reisz 1999; Hill 2005; Rieppel 2001, 2007 of the skull, and the absence of dentition on the trans- and references therein). The group as a whole displays verse flange of the pterygoid (e.g. Modesto & Damiani awidevarietyofbodyshapesandsizes,exemplifiedby 2007; Sail¨ a2008).Procolophonoidsaretheonlypararep-¨ the gracile and fully aquatic mesosaurs (e.g. Oelofsen & tiles known to have survived the end-Permian mass extinc- Araujo´ 1983), the tiny bipedal bolosaurs (e.g. Berman et al. tion event, and have been widely discussed in the context 2000) and the bulky, often heavily armoured pareiasaurs of survivorship rate analyses across the Permian-Triassic (e.g. Lee 1997b). In recent years taxonomic reviews and boundary (examples of similar analyses applied to other systematic efforts have improved our knowledge of these tetrapod groups include: Modesto & Rybczynski 2000; amniotes. As a result, detailed analyses of the intrinsic and Spencer & Benton 2000; Modesto et al.2001,2002,2003; external relationships of parareptiles as a whole, and of Benton et al.2004;Cisneroset al.2005;Rutaet al.2007; several constituent clades in this group, are now available Cisneros 2008a; Frobisch¨ 2008; Ruta & Benton 2008). (e.g. Rieppel & deBraga 1996; deBraga & Rieppel 1997; Procolophonoids are both morphologically and ecolog- Modesto et al.2003;Cisneroset al.2004;Pineiro˜ et al. ically diverse (Cisneros 2006, 2008a). They have been ∗Corresponding author. Email: [email protected] ISSN 1477-2019 print / 1478-0941 online Copyright C 2010 The Natural History Museum " DOI: 10.1080/14772019.2010.491986 http://www.informaworld.com 608 J. C. Cisneros and M. Ruta recorded from all continents and encompass some forty These two data categories should not be regarded as mutu- species with a broad range of morphofunctional adapta- ally exclusive or incompatible, as each is suited to specific tions. For example, their dentition attests to a wide spectrum targets and each presents advantages and limitations (see of feeding specializations in the herbivory guild (e.g. Gow comments in Wills et al.1994).Bearingthisinmind, 1977; Reisz & Sues 2000; Reisz 2006; Cisneros 2008a). an empirical (sensu McGhee 1999) morphospace based Numerous taxa exhibit impressive cranial sculptures in the upon cladistic data (i.e. a character-state multidimensional form of spikes and/or frills, the functional role of which may space), derived from a multivariate treatment of intertaxon have been related to protection, display and/or recognition distances, effectively depicts the morphological variety of (e.g. Cisneros 2008c). taxa under study. Procolophonoids provide an interesting case study for In the rest of this paper we use the largest clade elucidating macroevolutionary trends, patterns of morpho- within Procolophonoidea, the family Procolophonidae (see logical transformation, and rates of character change in Cisneros 2008a, b; Tsuji & Muller¨ 2009), as the basis for tetrapods in the aftermath of the end-Permian extinction a disparity analysis (the first of its kind for parareptiles) (Benton et al.2004).Thegroupisamenabletothese using cladistic characters. We hope to pave the way for kinds of analyses because the study of its relationships is similar analyses in other parareptile groups, and to encour- significantly advanced over those of other parareptiles, and age exploration of other data sources (e.g. landmark-based thus permits the exploration of morphological variety using geometric morphometrics). discrete character sets (e.g. Foote 1991, 1992; Wills et al. 1994; Roy & Foote 1997; Wills 1998; Brusatte et al. 2008; Aims Ruta 2009; Young et al.2010). In this paper we present a detailed analysis of morpholog- In recent years there has been a significant resurgence ical disparity in procolophonids. Our investigation draws of interest in studies of biodiversity and ecological change from a recently published phylogenetic analysis (Cisneros through time, as well as in the impact of large-scale extinc- 2008a, b) of the group, and uses the cladistic data matrix tions on diversification, especially in terrestrial environ- of that study as the starting point for a quantitative treat- ments. Several of these studies include analyses of morpho- ment of the degree of morphological similarity among taxa logical diversity (hereafter, disparity) which have found and their distribution in morphospace (more accurately, a broad applications in both phylogenetic and macroecologi- character-state space; e.g. Foote 1991, 1992; Wills et al. cal frameworks (e.g. Wills et al.1994;Wagner1995,1997; 1994; Roy & Foote 1997; McGhee 1999; McGowan 2007; Foote 1997; Roy & Foote 1997; Wills 1998; Ciampaglio Ruta 2009; Young et al.2010). et al.2001;Ciampaglio2002;Villier&Korn2004;Wesley- Our goals are: (1) to explore patterns of morphospace Hunt 2005; Stayton 2006; Botha & Angielczyk 2007; occupation for procolophonids as a whole and for McGowan 2007; Brusatte et al.2008;Sidlauskas2008; constituent groups in this clade; (2) to quantify disparity Benton 2009; Ruta 2009; Young et al.2010;foradditional for different sets of taxa grouped by stratigraphic age, remarks, see also McGhee 1999; Eble 2000; Erwin 2007, ecological adaptations, phenotypical resemblance, and 2008). phylogenetic relatedness; and (3) to reconstruct a palaeo- Here we use cladistic characters as a basis for biogeographical scenario, including inferred modalities disparity calculations and to construct a character-state
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