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Phylogeny of the Carnivoramorpha: the Impact of Postcranial Characters Michelle Spaulding a B D & John J

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Phylogeny of the Carnivoramorpha: the Impact of Postcranial Characters Michelle Spaulding a B D & John J This article was downloaded by: [American Museum of Natural History] On: 01 February 2013, At: 10:50 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Journal of Systematic Palaeontology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tjsp20 Phylogeny of the Carnivoramorpha: The impact of postcranial characters Michelle Spaulding a b d & John J. Flynn b c a Department of Earth & Environmental Sciences, Columbia University, New York, NY, 10027, USA b Division of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024, USA c Richard Gilder Graduate School, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024, USA d Section of Mammals, Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, Pennsylvania, 15206, USA Version of record first published: 25 Apr 2012. To cite this article: Michelle Spaulding & John J. Flynn (2012): Phylogeny of the Carnivoramorpha: The impact of postcranial characters, Journal of Systematic Palaeontology, 10:4, 653-677 To link to this article: http://dx.doi.org/10.1080/14772019.2011.630681 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, 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. 10, Issue 4, December 2012, 653–677 Phylogeny of the Carnivoramorpha: The impact of postcranial characters Michelle Spauldinga,b,d∗ and John J. Flynnb,c aDepartment of Earth & Environmental Sciences, Columbia University, New York, NY 10027, USA; bDivision of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA; cRichard Gilder Graduate School, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA; dSection of Mammals, Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, Pennsylvania 15206, USA (Received 30 August 2010; accepted 24 May 2011; printed 5 December 2012) Previous analyses of the mammalian clade Carnivoramorpha have tended to sample only cranial and dental character systems, particularly those that included early Cenozoic taxa, Viverravidae and basal Carnivoraformes (the latter being a paraphyletic array of taxa traditionally referred to as the ‘Miacidae’). In this study we add more than 100 postcranial characters to an existing dataset dominated by cranio-dental characters. The addition of the new characters permits the inclusion of a large number of basal carnivoraforms, known solely or predominantly from postcranial characters, that previously would have been ‘unplaceable’ in a phylogenetic analysis. The resultant phylogeny recovers most of the same clades identified in previous studies, but resolves some relationships differently within the basal carnivoraforms. A novel (unnamed) monophyletic subclade of the Carnivoraformes is recovered, supported in part by characters from both the prior and new datasets. The inclusion of a substantial suite of postcranial characters expands the ability to assess the relationships of basal carnivoramorphan taxa, and permits the inclusion of many taxa represented only by incomplete material. Keywords: Carnivoramorpha; Carnivora; postcrania; Eocene; Carnivoraformes; Viverravidae Introduction mammals, seeking to determine if there was a significantly different level of homoplasy in cranial, dental or postcra- Carnivoramorphan phylogeny has been the focus of a nial character sets. They found, measured by the consistency number of recent morphological cladistic studies (e.g. Wyss index, that there was no statistically significant difference in & Flynn 1993; Wesley-Hunt & Flynn 2005). Those attempts the resolving power of characters from any of these arbitrary to reconstruct relationships of taxa within this clade relied partitions. In addition, studies with a functional morphol- primarily upon cranial and dental characters, to near total ogy emphasis (e.g. Sargis 2002; Young2008) have routinely exclusion of the postcranial skeleton (only five postcranial found that the impact of phylogeny cannot be discounted characters were included in both cited studies). Postcranial when examining postcranial adaptations. characters could be an important source of phylogenetic Several previous studies particularly sought to clarify information, however, as they also should reflect synapo- the relationships between carnivoramorphan taxa from the morphies due to shared ancestry. Serious undersampling Palaeocene and Eocene and crown clade Carnivora (e.g. of postcranial characters in carnivoramorphan phylogeny Flynn & Galiano 1982; Flynn et al. 1988; Wyss & Flynn is especially striking in light of several studies of other 1993; Wesley-Hunt & Flynn 2005 – see this latter paper for mammalian orders that include substantial data from the an in-depth review of prior studies). These early Cenozoic Downloaded by [American Museum of Natural History] at 10:50 01 February 2013 postcranium, such as for Primates (Bloch et al. 2007), fossil taxa generally have been reconstructed to be either Artiodactyla (including cetaceans; Spaulding et al. 2009) members of the monophyletic clade Viverravidae or basal and Pholidota (Gaudin et al. 2009). Potential explanations Carnivoraformes (Carnivoraformes name following Flynn for this lack of inclusion are the difficulty of associating et al. 2010). The Viverravidae are known from the early postcrania and craniodental elements in disarticulated spec- Palaeocene to late Eocene, and have been firmly estab- imens, and longstanding assumptions (untested) that the lished as a monophyletic clade that is the nearest outgroup skeleton of early carnivoramorphans resembled a presumed to the rest of the Carnivoramorpha (Wesley-Hunt & Flynn generalized placental condition and many specializations 2005; Spaulding & Flynn 2009). Basal Carnivoraformes observed reflected function (such as locomotion) rather are known from the late Palaeocene to late Eocene. This than phylogeny. On this latter point, Sanchez-Villagra & array of early carnivoraform taxa, previously referred to as Williams (1998) examined a large number of datasets for the ‘Miacidae’, has been documented to be paraphyletic ∗Corresponding author. Email: [email protected] ISSN 1477-2019 print / 1478-0941 online Copyright C 2012 The Natural History Museum http://dx.doi.org/10.1080/14772019.2011.630681 http://www.tandfonline.com 654 M. Spaulding and J. J. Flynn in a number of the studies cited above, and their potential associated with no postcranial material, and one preserves monophyly was firmly refuted in the latest comprehensive little if any basicranium. morphological phylogeny (Wesley-Hunt & Flynn 2005). In this study we demonstrate the utility of postcranial However, a recent cladistic study of this diverse suite of characters in resolving relationships among the Carnivo- basal Carnivoraformes has shown that our ability to resolve ramorpha. To accomplish this goal, a postcranial data their interrelationships is limited in the absence of postcra- matrix was constructed and combined with the most nial characters (Spaulding & Flynn 2009). When the topol- comprehensive existing craniodental matrix (Wesley-Hunt ogy derived from the character matrix of Wesley-Hunt & & Flynn 2005). All taxa previously included in the cranio- Flynn (2005; emphasizing basicranial and dental charac- dental matrix analyses were included, and other taxa repre- ters) is compared with subsequent studies (e.g. Wesley- sented by scant basicranial material were added. Several Hunt & Werdelin 2005; Polly et al. 2006) using a similar analyses of this combined matrix were then performed, each character set but incorporating additional taxa with rela- utilizing different character subsets and sets of taxa. Inclu- tively complete dental and basicranial material, there is sion of postcranial characters in the combined matrix not little if any change in topology from the earlier analysis. only serves to provide more character information for taxa However, when additional taxa lacking complete dental and already included in prior analyses, thus providing an addi- basicranial material, such as “Miacis” uintensis (Spauld- tional test of previously proposed hypotheses of relation- ing & Flynn 2009) and Dawsyonicyon isami (Spauld- ships, but also permits the expansion of taxon sampling. ing et al. 2010), are added to the analysis, the topology This substantially enhanced sampling of characters and among basal taxa becomes more poorly resolved and less taxa, in turn, should give us the ability to erect more compre- stable, indicated by an increase in the number of poly-
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