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Research Article

Total evidence phylogenetic analysis of Leiosauridae (Squamata) with focus on the ‘para-anoles’ and description of a new Urostrophus species from the Bolivian Andes

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Abstract

Only since the late 1980s have lizard specialists begun incorporating phylogenetic information to the classification of leiosaurids. These are among the most enigmatic South American lizards, with 35 currently recognised species arranged in six genera. To evaluate their relationships, we assembled a data set based on published and novel information containing 135 morphological characters and 11,235 DNA base pairs (14 loci) for up to 52 terminals, including outgroups. We performed 13 analyses based on different optimality criteria (parsimony, maximum likelihood, and coalescent approach) and indel coding strategies (gaps coded as missing data or as a fifth state). We also evaluated the impact of phenotypic evidence on the phylogeny of leiosaurids, and investigated the level of informativeness and potential sources of topological conflict using a Quartet Sampling analysis. Phylogenetic analyses supported the monophyly of major leiosaurid clades, including subfamilies Enyaliinae and Leiosaurinae, and the ‘para-anoles’ clade (Urostrophus + Anisolepis). However, Urostrophus was inferred as paraphyletic, with U. vautieri forming a clade with Anisolepis. These relationships motivate the synonymization of Anisolepis Boulenger, Citation1885 with Urostrophus Duméril & Bibron, 1837. The monophyly of Pristidactylus and, surprisingly, of Enyalius, were also challenged, underscoring the need for additional phylogenetic work. Finally, we described Urostrophus chungarae sp. nov. from the Torotoro National Park, Potosí, Bolivia. This apparently rare leiosaurid was inferred as sister to U. gallardoi, a Chacoan species distributed in northern Argentina and southeastern Bolivia. We conclude with a revised classification of Leiosauridae and by pointing out a number of unresolved issues that require further systematic investigation.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:F60EB6CB-62B2-4EE3-B1A8-BEE3B2ED7A65

Acknowledgements

In memory of Richard E. Etheridge for his infinite enthusiasm and invaluable teaching about leiosaurids. We express our sincerest gratitude to Eliana Lizarraga, Gabriel Callapa, Arturo Muñoz, Lidia Meneses, and Teresa Camacho, from the Museo de Historia Natural Alcide d’Orbigny (MNHNC), for the advice and logistic support provided during fieldwork in Bolivia in 2013, and/or during analysis of specimens in 2013 and 2017. We thank Kathia Rivero, Edson Cortez Cuellar, and Libertad Ramos from the Museo de Historia Natural Noel Kempff Mercado (MNK), for the valuable assistance during analysis of specimens in 2013 and 2017. We are in debt to Felipe Toledo from Universidade de Campinas (UNICAMP–ZUEC) for facilitating the loan of specimens of Urostrophus vautieri for morphological and hemipenial analysis. We thank the Bolivian Ministerio de Medio Ambiente y Agua (MMAyA) for collection permits and the administration of Torotoro National Park for support during our visit in 2013. We also thank the park rangers Eduardo Lujo, Felix Mamani, Geronimo Juchazara, Justino Kamaqui, and Juvenal Choque for guiding us inside the area of the park. Renato Gaiga (Biotropica Consultoria), Ivan Prates (U-M), Diego Emmanuel Oscar (independent photographer), Patricia Mancilla (independent photographer), Cristian Abdala (UEL-CONICET), Arturo Muñoz (MNHNC), Lucindo Gonzales (MNK), Gabriel Callapa (MNHNC), Rafael Paradero (UFMT), and Welington Coelho (UnB) made available to us important field records and/or photographs included/analysed in this study. AL thanks Santiago Nenda (MACN), Julián Faivovich (MACN), Soledad Palomas (UNNE), Fernando Lobo (MCN), José Artigas (MZUC), Sonia Kretzschmar (FML), and Gustavo Scrocchi (FML) for assisting while examining collections under their care. We thank Sebastián Quinteros (UNSa) and Pablo Goloboff (UEL) for advice on the treatment of continuous morphological characters. We thank Maria del Rosario Castañeda (Universidad ICESI – Cali) for suggestions on PCR protocols for amplification of RAG1 fragments, and Jose Maldonado and Carlee Painter (UTA) for performing the PCR. We are grateful to Adam Leaché and Hayden Davis (UW), Mark Wilkinson (BMNH), Rachel Montesinos (UFMG), and Danilo Giacometti (Brock University) for critically revising (in part or completely) the first draft of the manuscript.

Disclosure statement

The authors report there are no competing interests to declare.

Supplemental material

Supplemental material for this article can be accessed here: http://dx.doi.org/10.1080/14772000.2023.2200306.

        Associate Editor: Dr David Gower

Additional information

Funding

This work was supported by the Brazilian National Council for Scientific and Technological Development (CNPq) under grants #200798/2010-3 and #302864/2020-2; the São Paulo Research Foundation (FAPESP) under grants #2016/08249-6, #2017/20235-3, and #2011/50146-6; the Argentinian National Scientific and Technical Research Council (FONCyT–ANPCyT) under grant #PICT 2017-0553; the Brazilian Research Support Foundation of the Federal District (FAPDF) under grant #0193.001637/2017; the Rufford Foundation under grant #2013/13081-2; besides receiving additional financial aid from the Explorers Club, the Andrew Sabin Family Foundation, and the American Museum of Natural History (RGGS/AMNH).
The Explorers Club, Andrew Sabin Family Foundation, the American Museum of Natural History (RGGS/AMNH), and the São Paulo Research Foundation (FAPESP) provided financial support to ALGC for fieldwork and/or analysis of museum specimens in Bolivia. ALGC was also supported by the Brazilian National Council for Scientific and Technological Development (CNPq) (#200798/2010-3) during his Ph.D. at the AMNH, and a postdoctoral fellowship from FAPESP (#2016/08249-6; #2017/20235-3). MTR thanks FAPESP (#2011/50146-6) and CNPq (#302864/2020-2). AL is a postdoctoral fellow of the Argentinian National Scientific and Technical Research Council (FONCyT–ANPCyT; #PICT 2017-0553). LRR thanks the Rufford Foundation for financing the project ‘Conservation of threatened and endemic reptiles of the inter-Andean dry valleys and highlands of Bolivia’ (#2013/13081-2). JK thanks the Brazilian Research Support Foundation of the Federal District (FAPDF) for financial support (#0193.001637/2017).

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