Acta Herpetologica 9(2): 249-252, 2014 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-14562 No evidence for conspecificity between two high Andes Liolaemus lizards (Squamata: Liolaemidae) Daniel Pincheira-Donoso Laboratory of Evolutionary Ecology of Adaptations, School of Life Sciences, University of Lincoln, Brayford Campus, Lincoln, LN6 7DL, Lincolnshire, United Kingdom. E-mail: DPincheiraDonoso@lincoln.ac.uk Submitted on 2014, 26th May; revised on 2014, 10th September; accepted on 2014, 11th October Editor: Aaron M. Bauer Abstract. The remarkable taxonomic, ecological and geographic diversity achieved by the South American lizard genus Liolaemus has inspired persistent debate about species boundaries and the reliability of phenotypic predictors/ indicators of reproductive isolation between species (i.e., signatures of speciation). Many aspects of these debates remain unsettled and part of the diversity of the genus remains under controversy. Factors such as small samples, or lack of molecular data to quantify genetic differences between species can be regarded as legitimate limitations on the ability to draw definite taxonomic conclusions. However, conclusions drawn from careless and negligent observations should be taken with high degree of caution. A recent paper offers a clear example of this latter scenario, in which it is suggested that Liolaemus filiorum Pincheira-Donoso and Ramirez, 2005 is a synonym of Liolaemus puritamensis Nuñez and Fox, 1989 based on qualitative analyses of the “holotype” of the former which is, in fact, not the holotype of this taxon, but rather one of the paratypes of the latter species. Editors and referees should play a central role in preventing publication of studies of this nature. Keywords. Liolaemus, speciation, synonymy, taxonomy, integrative taxonomy. The iguanian genus Liolaemus is one of the most spe- cies-rich groups of living vertebrates (Pincheira-Donoso et al., 2013a). The prolific adaptive radiation undergone by this lineage has resulted in 240+ species adapted to a remarkable diversity of ecological conditions, encompass- ing all climatic extremes found in central and southern South America, including deserts, tropical environments, and some of the highest elevations/latitudes recorded among reptiles globally (Morando et al., 2003; Espinoza et al., 2004; Schulte et al., 2004; Pincheira-Donoso et al., 2008, 2013b; Labra et al., 2009; Pincheira-Donoso, 2011; Pincheira-Donoso and Tregenza, 2011; Meiri et al., 2013). Not surprisingly, the causes and consequences, and the genetic and phenotypic expressions behind such diversity have been the focus of persistent debate and controver- sy (Morando et al., 2003, 2004; Pincheira-Donoso and Nuñez, 2005; Avila et al., 2006; Labra, 2011; Pinchei- ra-Donoso, 2012). Opinions remain divergent, and debates remain unsettled. In recent years, the development of the ‘integrative taxonomy’ approach has made explicit the need for tax- onomic conclusions to be drawn from quantitative and multivariate analyses of the various components involved in the process of species formations, i.e., the search for signatures of speciation (Padial et al., 2009, 2010). This approach adds scientific objectivity to taxonomic conclu- sions, and has in fact increasingly been adopted by liolae- mid researchers (Avila et al., 2006; Morando et al., 2007; Pincheira-Donoso et al., 2007; Breitman et al., 2011a, b; Morando et al., 2013; Scolaro et al., 2013). However, a recent study (Troncoso-Palacio, 2014) that proposed a conclusion with major implications (conspecificity between two species) not only employs a purely typolog- ical and qualitative analytical approach, but is based on a fundamental error. In this study, Troncoso-Palacio (2014) concludes that Liolaemus puritamensis Nuñez and Fox, 1989 and L. filiorum Pincheira-Donoso and Ramirez, 2005 are the same species (given the temporal priority of 250 Daniel Pincheira-Donoso L. puritamensis, the latter taxon is treated as a subjective junior synonym of the former). This conclusion is based on comparisons between three specimens, two L. purita- mensis and (as claimed by the author) the “holotype” of L. filiorum. Although the lack of a quantitative analytical approach and the extremely limited sample sizes may be viewed as inappropriate to support the hypothesis of con- specificity between these taxa, the very basic flaw that invalidates the conclusions of Troncoso-Palacio (2014) is that this “holotype” is not the holotype of L. filiorum but, in fact, one of the paratypes of L. puritamensis itself (Fig. 1). This specimen was recently transferred to the Museo Nacional de Historia Natural de Chile collection from another collection and erroneously given the same num- ber, MNHN-3829, as the L. filiorum holotype, although the data accompanying the specimen clearly identify it as a paratype of L. puritamensis. By focusing on the museum collection number of the specimen rather than on details of locality, date and collectors provided in the original description of L. filiorum, Troncoso-Palacio compared specimens in the same type series and not surprisingly concluded that these specimens were conspecific. Thus, no evidence exists to conclude that these two Liolaemus are the same species. Here, I provide images of the real holo- Fig. 1. Liolaemus from the High Andes of Chile. Liolaemus filiorum: Dorsal views of the holotype (A), paratype (B), and two living adult male specimens (C, D) from the type locality, Cerro Las Papas, Chile. Liolaemus puritamensis: A living specimen of from Vilaco, Chile (E). 251No conspecificity in Liolaemus type (MNHN-3829) and paratype (CHDPD-01069) of L. filiorum, and pictures of both species in life (Fig. 1). Taxonomy has seen a tendency to be treated as a marginal discipline, especially in high-impact scientific journals, and has been considered the ‘Cinderella’ of biol- ogy (e.g., Padial and De la Riva, 2007). While this view disregards the overwhelmingly vital role of taxonomic research as the basis for understanding biodiversity, and thus for areas such as ecology and evolution (see Isaac et al., 2004, for a discussion), it is essential that taxonomists conduct this science with due care and scientific rigor. Taxonomy is a science and, as such, it depends on the use of scientific approaches to thoroughly and quantitatively test hypotheses (e.g., hypotheses of speciation events as the basis for recognizing different species, or evidence that the genomes of two species fail to show signals of speciation events, which can include phenotypic or genet- ic signals). In this context, it is crucial for editors, refer- ees and colleagues to identify studies that do not employ such standards before they are approved for publication. ACKNOWLEDGEMENTS The author thanks an anonymous referee for con- structive comments on an earlier version of this man- uscript. Also, thanks to the financial support provided by a University of Lincoln Research Investment Fund Grant (RIF). REFERENCES Avila, L.J., Morando, M., Sites, J.W. (2006): Congener- ic phylogeography: hypothesizing species limits and evolutionary processes in Patagonian lizards of the Liolaemus boulengeri group (Squamata : Liolaemini). Biol. J. Linnean Soc. 89: 241-275. Breitman, M.F., Avila, L.J., Sites, J.W., Morando, M. (2011a): Lizards from the end of the world: phyloge- netic relationships of the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemini). Mol. Phylo- genet. Evol. 59: 364-376. Breitman, M.F., Perez, C.H.F., Parra, M., Morando, M., Sites, J.W., Avila, L.J. (2011b): New species of lizard from the magellanicus clade of the Liolaemus lineo- maculatus section (Squamata: Iguania: Liolaemidae) from southern Patagonia. Zootaxa 3123: 32-48. Espinoza, R.E., Wiens, J.J., Tracy, C.R. (2004): Recurrent evolution of herbivory in small, cold-climate lizards: breaking the ecophysiological rules of reptilian her- bivory. Proc. Natl. Acad. Sci. USA 101: 16819-16824. Isaac, N.J.B., Mallet, J., Mace, G.M. (2004): Taxonomic inflation: Its influence on macroecology and conser- vation. Trends Ecol. Evol. 19: 464-469. Labra, A. (2011): Chemical stimuli and species recogni- tion in Liolaemus lizards. J. Zool. 285: 215-221. Labra, A., Pienaar, J., Hansen, T.F. (2009): Evolution of thermal physiology in Liolaemus lizards: adaptation, phylogenetic inertia, and niche tracking. Am. Nat. 174: 204-220. Meiri, S., Bauer, A.M., Chirio, L., Colli, G.R., Das, I., Doan, T.M., Feldman, A., Castro-Herrera, F., Novosolov, M., Pafilis, P., Pincheira-Donoso, D., Powney, G., Torres-Carvajal, O., Uetz, P., Van Damme, R. (2013): Are lizards feeling the heat? A tale of ecology and evolution under two temperatures. Global Ecol. Biogeogr. 22: 834-845. Morando, M., Avila, L.J., Sites, J.W. (2003): Sampling strategies for delimiting species: genes, individuals, and populations in the Liolaemus elongatus-kriegi Complex (Squamata: Liolaemidae) in Andean-Patago- nian South America. Syst. Biol. 52: 159-185. Morando, M., Avila, L.J., Baker, J., Sites, J.W. (2004): Phy- logeny and phylogeography of the Liolaemus darwinii complex (Squamata: Liolaemidae): Evidence for intro- gression and incomplete lineage sorting. Evolution 58: 842-861. Morando, M., Avila, L.J., Turner, C.R., Sites, J.W. (2007): Molecular evidence for a species complex in the pata- gonian lizard Liolaemus bibronii and phylogeography of the closely related Liolaemus gracilis (Squamata: Liolaemini). Mol. Phylogenet. Evol. 43: 952-973. Morando, M., Avila, L.J., Perez, C.H.F., Hawkins, M.A., Sites, J.W. (2013): A molecular phylogeny of the lizard genus Phymaturus (Squamata, Liolaemini): implica- tions for species diversity and historical biogeography of southern South America. Mol. Phylogenet. Evol. 66: 694-714. Nuñez, H., Fox, S.F. (1989): Liolaemus puritamensis, a new species of iguanid lizard previously confused with Liolaemus multiformis (Squamata: Iguanidae). Copeia 1989: 456-460. Padial, J.M., De la Riva, I. (2007): Taxonomy, the Cinder- ella of science, hidden by its evolutionary stepsister. Zootaxa 1577: 1-2. Padial, J.M., Miralles, A., De la Riva, I., Vences, M. (2010): The integrative future of taxonomy. Front. Zool. 7: 16. Padial, J.M., Castroviejo-Fisher, S., Kohler, J., Vila, C., Chaparro, J.C., De la Riva, I. (2009): Deciphering the products of evolution at the species level: the need for an integrative taxonomy. Zool. Scripta 38: 431-447. Pincheira-Donoso, D. (2011): Predictable variation of range-sizes across an extreme environmental gradient 252 Daniel Pincheira-Donoso in a lizard adaptive radiation: evolutionary and eco- logical inferences. PLoS One 6: e28942. Pincheira-Donoso, D. (2012): Cautionary comments on the influence of chemical-based interactions as poten- tial drivers of sexual speciation in Liolaemus lizards. J. Zool. 288: 231-233. Pincheira-Donoso, D., Nuñez, H. (2005): Las especies chilenas del género Liolaemus. Taxonomía, sistemáti- ca y evolución. Publ. Ocas. Mus. Nac. Hist. Nat. Chile, 59: 1-487. Pincheira-Donoso, D., Ramirez, G.M. (2005): Despla- zamiento de caracteres como evidencias de un posi- ble caso de especiación simpátrica entre dos Liolae- mus del grupo jamesi en la Provincia de El Loa, con la descripción de una nueva especie. In: Fauna del Altiplano y Desierto de Atacama. Vertebrados de la Provincia de El Loa, pp. 350-365. Ramírez, G.M., Pincheira-Donoso, D., Eds, Phrynosaura Ediciones, Calama, Chile. Pincheira-Donoso, D., Tregenza, T. (2011): Fecundity selection and the evolution of reproductive output and sex-specific body size in the Liolaemus lizard adaptive radiation. Evol. Biol. 38: 197-207. Pincheira-Donoso, D., Scolaro, J.A., Schulte, J.A. (2007): The limits of polymorphism in Liolaemus rothi: molecular and phenotypic evidence for a new spe- cies of the Liolaemus boulengeri clade (Iguanidae, Liolaemini) from boreal Patagonia of Chile. Zootaxa 1452: 25-42. Pincheira-Donoso, D., Hodgson, D.J., Tregenza, T. (2008): The evolution of body size under environmental gra- dients in ectotherms: why should Bergmann’s rule apply to lizards? BMC Evol. Biol. 8: 68. Pincheira-Donoso, D., Bauer, A.M., Meiri, S., Uetz, P. (2013a): Global taxonomic diversity of living reptiles. PLoS One 8: e59741. Pincheira-Donoso, D., Tregenza, T., Witt, M.J., Hodgson, D.J. (2013b): The evolution of viviparity opens oppor- tunities for lizard radiation but drives it into a climat- ic cul-de-sac. Global Ecol. Biogeogr. 22: 857-867. Schulte, J.A., Losos, J.B., Cruz, F.B., Nuñez, H. (2004): The relationship between morphology, escape behaviour and microhabitat occupation in the lizard clade Liolaemus (Iguanidae: Tropidurinae: Liolaemi- ni). J. Evol. Biol. 17: 408-420. Scolaro, J.A., Jara, M., Pincheira-Donoso, D. (2013): The sexual signals of speciation? A new sexually dimor- phic Phymaturus species of the patagonicus clade from Patagonia Argentina. Zootaxa 3722: 317-332. Troncoso-Palacio, J. (2014): Revisión del estatus taxonómico de Liolaemus filiorum Pincheira-Donoso y Ramírez, 2005 (Iguania: Liolaemidae). Cuad. Herpe- tol. 28: 1-7.