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The Identities of Liolaemus Signifer(Duméril & Bibron 1837)

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The Identities of Liolaemus Signifer(Duméril & Bibron 1837) Trabajo Cuad. herpetol. 35 (Supl. 1): 111-167 (2021) The identities of Liolaemus signifer (Duméril & Bibron 1837), L. pantherinus Pellegrin 1909, L. schmidti (Marx 1960), and L. erroneus (Núñez & Yáñez 1984 “1983-1984”) (Squamata: Liolaemidae) Roberto Langstroth Plotkin Colección Boliviana de Fauna, Calle 27 Cota Cota, La Paz, Bolivia. Recibido: 27 Abril 2020 ABSTRACT Revisado: 17 Julio 2020 Based on detailed examination of the literature, visits to collections and the field, correspondence Aceptado: 24 Agosto 2020 with specialists and curators, study of detailed photography of living and preserved specimens, Editor Asociado: A. S Quinteros and statistical analysis of meristic data, I propose a number of taxonomic and nomenclatural changes for species of the Liolaemus montanus group from the Andes of Bolivia, southern Peru, northern Chile and northwestern Argentina. Liolaemus signifer (Duméril & Bibron 1837) is de- doi: 10.31017/CdH.2020.(2020-094) signated a nomen dubium and Liolaemus multiformis (Cope 1875) and Liolaemus lenzi Boettger 1891 are revalidated. Liolaemus variabilis Pellegrin 1909 and Liolaemus bolivianus Pellegrin 1909 are placed in the synonymy of L. lenzi. Liolaemus pleopholis Laurent 1998 is considered a species inquirenda allied with L. lenzi. Liolaemus annectens Boulenger 1901 is considered a valid species and Liolaemus tropidonotus Boulenger 1902 is considered a junior synonym of L. multiformis. Liolaemus pantherinus Pellegrin 1909 is recognized as a valid species that includes L. annectens orientalis Müller 1924 and L. multiformis simonsii—Burt & Burt 1931 are included in its synonymy. Liolaemus islugensis Ortiz & Marquet 1987 is removed from the synonymy of L. pantherinus and is placed in the synonymy of L. schmidti (Marx 1960). Liolaemus erguetae Laurent 1995 and L. molinai Valladares et al. 2002 are placed in the synonymy of L. erroneus (Núñez & Yáñez 1984 “1983–1994”). Key Words: Lizards; Taxonomy; Nomenclature; Synonyms; Andes. RESUMEN Basado en la revisión detallada de la literatura, visitas a colecciones y el campo, correspon- dencia con especialistas y curadores, estudio de fotografías detalladas de ejemplares vivos y preservados y análisis estadístico de datos merísticos, se propone varios cambios taxonómicos y nomenclaturales para especies del grupo Liolaemus montanus de los Andes de Bolivia, el sur del Perú, norte de Chile y noroeste de Argentina. Liolaemus signifer (Duméril & Bibron 1937) se designa nomen dubium y se revalidan Liolaemus multiformis (Cope 1875) y Liolaemus lenzi Boettger 1891. Liolaemus variabilis Pellegrin 1909 y Liolaemus bolivianus Pellegrin 1909 se consideran sinónimos júniores de L. lenzi. Liolaemus pleopholis Laurent 1998 se considera una species inquirenda aliada a L. lenzi. Liolaemus annectens Boulenger 1901 se considera especie válida y Liolaemus tropidonotus Boulenger 1902 se considera sinónimo júnior de L. multifor- mis. Liolaemus pantherinus Pellegrin 1909 se reconoce como especie válida, la cual incluye L. annectens orientalis Müller 1924 y L. multiformis simonsii—Burt & Burt 1931 en su sinonimia. Liolaemus islugensis Ortiz & Marquet 1987 se transfiere de la sinonimia de L. pantherinus a la sinónima de L. schmidti (Marx 1960). Liolaemus erguetae Laurent 1995 y L. molinai Valladares et al. 2002 se consideran sinónimos júniores de L. erroneus (Núñez & Yáñez 1984 “1983–1994”). Palabras claves: Lagartijas; Taxonomía; Nomenclatura; Sinónimos; Andes. Introduction The lizard genus Liolaemus Wiegmann 1834 is of which over 270 are recognized as valid (Abdala highly diverse with more than 350 published na- et al., this volume). In addition, there is a growing mes (including species, subspecies, and varieties), number of “candidate species” identified by mole- Author for correspondence: [email protected] 111 R. Langstroth Plotkin - Taxonomic and nomenclatural changes in Liolaemus montanus group cular, morphological, and combined phylogenetic Materials and methods analyses (for example, see Abdala et al., 2020; Aguilar-Puntriano et al., 2018). While modern This paper is the result of an analysis of the relevant phylogenetic assessments have clearly demonstrated body of literature (for example, Duméril and Bibron, the monophyly of Liolaemus, the genus is generally 1837; Bell, 1843; Cope, 1875; Boettger, 1891; Pelle- recognized as included two major clades: Liolaemus grin, 1909; Marx, 1960; Cei et al., 1980; Laurent, sensu stricto (the “chileno” group of Laurent, 1983) 1982, 1984a, 1992, 1995); Ortiz and Marquet, 1987; and Eulaemus Girard 1858 (the “argentino” group of Núñez and Yáñez, 1984 “1983-1984”; Valladares et Laurent, 1983). Within Eulaemus, the monophyletic al., 2002; and Aguilar-Puntriano et al., 2018), review L. montanus group (sensu Etheridge, 1995; Abdala of itineraries of collectors (d’Orbigny 1844, 1846; et al., 2020) includes some 60 species (Olave et al., Créqui Montfort and Sénéchal de la Grange, 1904; 2014; Aguilar et al., 2017a, Abdala et al., 2020) Neveu-Lemaire, 1904; Hellmayr, 1932), analysis which are largely restricted to high elevation “puna” of unpublished data and notes on type specimens environments of the Andes, with the exception of a generously provided by Richard E. Etheridge and few lowland species in the Pacific slope deserts and many other specialists and curators (see Acknowled- the extra-Andean Sierras of northern and central gements), examination of specimens, including Argentina (Abdala et al., 2020). As used here, the L. relevant type material during visits to collections montanus group is not equivalent to the L. monta- including the American Museum of Natural History nus section of Eulaemus recovered by Schulte et al. (AMNH), the Natural History Museum (BMNH), (2000), which includes a L. boulengeri series charac- the Colección Boliviana de Fauna (CBF), the Field terized by the presence of a patch of enlarged spinose Museum (FMNH), the Museum of Vertebrate Zoo- scales on the inner thigh; rather it is equivalent to logy (MVZ), the Colección de Flora y Fauna Patricio their L. montanus series. Sánchez Reyes, Pontificia Universidad Católica The earliest available name assigned to the de Chile (SSUC), and the Smithsonian National Liolaemus montanus group is Proctotretus signifer Museum of Natural History (USNM) and other Duméril and Bibron 1837, a species based on a material reviewed from detailed digital photographs single specimen that was lost sometime after it was of additional material from the Academy of Natural examined and redescribed by Bell (1843). Cei et al. Sciences of Philadelphia (ANSP), Carnegie Museum (1980) claimed to have found the lost holotype but of Natural History (CM), Fundación Miguel Lillo Laurent (1984a) noted significant discrepancies in (FML), Monte L. Bean Life Science Museum (BYU), the measurements reported by earlier authors and Museo de Historia Natural de la Universidad Na- those reported by Cei et al. (1980). Despite his cional de San Agustín (MUSA), Museo de Historia earlier misgivings, Laurent (1992) decided that L. Natural de la Universidad Nacional Mayor de San signifer is the senior synonym of number of species Marcos (MUSM), Museo de Zoología de la Univer- described from Peru and Bolivia: L. multiformis sidad de Concepción (MZUC), Museo Nacional de (Cope 1875), L. lenzi Boettger 1891, L. annectens Historia Nacional de Chile (MNHNCL), Museum Boulenger 1901, L. tropidonotus Boulenger 1902, L. für Naturkunde (ZMB), Museum of Comparative pantherinus Pellegrin 1909, L. variabilis Pellegrin Zoology (MCZ), Muséum National d’Histoire Natu- 1909, and L. bolivianus Pellegrin 1909. relle (MNHN), San Diego State University Museum The present paper will explore the taxonomic of Biodiversity (SDSU), Senckenberg Museum history of Liolaemus signifer and the identity of its (SMF), University of Kansas Biodiversity Institute purported discovered holotype and then clarify and Natural History Museum (KU), University of the status of the species included in its synonymy Michigan (UMMZ), Zoologische Museum Hamburg by Laurent (1992), with a special consideration of (ZMH), and Zoologische Staatssammlung München L. pantherinus and a suite of other misunderstood (ZSM), analysis of counts of dorsal scales between names with intertwined and convoluted histories: L. the occiput and the anterior margin of the thighs schmidti (Marx 1960), L. erroneus (Núñez & Yáñez (“DSOT”) of 190 specimens representing 14 species, 1984 “1983-1984”), L. islugensis Ortiz & Marquet and consideration of most complete phylogenetic 1987, L. erguetae Laurent 1995, and L. molinai Va- hypotheses and biogeographic patterns for the focal lladares, Etheridge, Schulte, Manríquez, & Spotorno species (Aguilar-Puntriano et al., 2018; Abdala et 2002. al., 2020). The complete list of material examined 112 Cuad. herpetol. 35 (Supl. 1): 111-167 (2021) is presented in Appendix 1. species previously described by Wiegmann (1834) DSOT counts are a frequently used meristic in the subgenus Tropidurus (Liolaemus) and eight character in descriptions and diagnoses of Liolae- new species based on material the Muséum Natio- mus species, referred to variably as “dorsals”, “pa- nal d’Histoire Naturelle (MNHN) in Paris. Their P. ravertebrals” or “middorsal scales” in the literature signifer was based on a single specimen attributed (Etheridge, 1995). The first dorsal row is treated as to the collections of d’Orbigny and having the sta- the first row of scales posterior to the enlarged scales ted locality of simply “Chili” (Duméril and Bibron, or plates (e.g., postoccipitals)
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