Abstract
Within the Liolaemus boulengeri group or ‘patch’ group, there is a monophyletic clade named Liolaemus fitzingerii group.This clade is composed of large Liolaemus individuals, mostly psammophilus, oviparous and mainly insectivorous, dis-tributed from central Río Negro and Neuquén to the south of Santa Cruz in Argentine Patagonia. We make a taxonomicand phylogenetic revision of the L. fitzingerii clade, describe two new species of the group, compare these results withphylogenetic hypotheses from the literature, and discuss the geographic distribution of species of the L. fitzingerii clade.The new species described here inhabit central and central-eastern Chubut Province, Argentina. These new species areeasily distinguished from the other species of the L. fitzingerii group by a combination of morphological characters. Thephylogenetic analyses support the description of these new species, because they are not closely related to the species withwhich they were previously confused. According to a new phylogenetic analysis of the L. fitzingerii clade, one of the newspecies, L. camarones sp. nov., has a relatively basal position within the group, whereas the other new species, L. shehuensp. nov., is sister taxon of L. chehuachekenk. With these two new species, the L. fitzingerii clade is now composed of nine species, all of which are distributed in Argentine Patagonia.References
Abdala, C.S. (2002) Nuevo Liolaemus (Iguania: Liolemidae) perteneciente al grupo boulengeri de la provincia de Neuquén, Argentina. Cuadernos de Herpetología, 16 (1), 3–13.
Abdala, C.S. (2003) Cuatro nuevas especies del género Liolaemus (Iguania: Liolaemidae), pertenecientes al grupo boulengeri, de la Patagonia Argentina. Cuadernos de Herpetología, 17 (1–2), 3–32.
Abdala, C.S. (2005a) Dos nuevas especies del género Liolaemus (Iguania: Liolaemidae) y redescripción de Liolaemus boulengeri (Koslowsky, 1898). Cuaderno de Herpetología, 19 (1), 3–33.
Abdala, C.S. (2005b) Sistemática y filogenia del grupo de L. boulengeri (Iguania: Liolemidae, Liolaemus) en base a caracteres morfológicos y moleculares. Tesis Doctoral, Universidad Nacional de Tucumán, 278 pp.
Abdala, C.S. (2007) Phylogeny of the boulengeri group (Iguania: Liolaemidae, Liolaemus) based on morphological and molecular characters. Zootaxa, 1538, 1–84.
Abdala, C.S. Quinteros, A. S. & Espinoza, R. E. (2008) Two new species of Liolaemus (Iguania: Liolaemidae) from the Puna of Northwestern Argentina. Herpetologica, 64, 458–471.
Abdala, C.S., Acosta, J.C., Cabrera, M.R. Villavicencio, H.J. & Marinero, J. (2009) A New Andean Liolaemus of the L. montanus Series (Squamata: Iguania: Liolaemidae) from Western Argentina. South American Journal of Herpetology, 4(2), 91–102.
Abdala, C.S. Quinteros, A.S., Scrocchi, G.J. & Stazzonelli, J.C. (2010) Three new species of the Liolaemus petrophilus group (Iguania: Liolaemidae) from Argentina. Cuadernos de Herpetología, 24 (1), 25–40.
Abdala, C.S. Quinteros, A.S., Arias, F., Portelli, S. & Pallavechino, A. (2011) A new species of the Liolaemus darwinii group (Iguania: Liolaemidae) from Salta Province, Argentina. Zootaxa, 2968, 26–38.
Abdala, V. Abdala, C. & Tulli, M.J. (2006) Muscular characters in the phylogeny of Liolaemus (Squamata: Iguania: Liolaemidae): a reappraisal. Zootaxa, 1205, 55–68.
Ávila, L.J. Morando, M. & Sites, J.W. Jr. (2006) Congeneric phylogeography: hypothesizing species limits and evolutionary processes in Patagonian lizards of the Liolaemus boulengeri group (Squamata: Liolaemini). Biological Journal of the Linnean Society, 89, 241–275.
Ávila, L.J., Perez, C.H.F., Morando, M. & Sites J.W. Jr. (2007) Geographic distribution: Liolaemus fitzingerii. Herpetological Review, 38 (3), 352–352.
Ávila, L.J., Morando, M. & Sites, J.W. Jr. (2008) New species of the iguanian lizard genus Liolaemus (Squamata, Iguania, Liolaemini) from central Patagonia, Argentina. Journal of Herpetology, 42, 186–196.
Ávila, L.J., Fluvio Perez, C. H. Morando, M. & Sister, J.W. JR. (2010) A new species of Liolaemus (Reptilia: Squamata) from southwestern Rio Negro Province, northern Patagonia, Argentina. Zootaxa, 2434, 47–59.
Breitman M.F., Ávila, L.J., Sites, J. W. & Morando, M. (2011) Lizards from the end of the world: Phylogenetic relationships of the Liolaemus lineomaculatus section (Squamata: Iguania: Liolaemini). Mol Phylogenet Evol., 59, 364–376.
Cabrera, A.L. & Willink, A. (1980) Biogeografía de América Latina. Secretaría General de la OEA. Prog. Reg. de Desarrollo Científico y Tecnológico, Washington D. C.
Cei, J.M. (1973) Herpetología patagónica. VI. Los Liolaemus del grupo fitzingerii en Santa Cruz y Chubut (Sauria, Iguanidae). Physis, 32, 447–458.
Cei, J.M. (1975) Liolaemus melanops (Burmeister) and the subspecific status of the Liolaemus fitzingerii group (Sauria, Iguanidae). Journal of Herpetology, 9, 217–222.
Cei, J.M. (1980) L’identité des syntypes de Proctotretus fitzingerii Duméril et Bibron, 1837. Bulletin Museum National d’Historie Naturelle (Paris), Ser. 4, A2, 317–320.
Cei, J.M. (1986) Reptiles del centro, centro-oeste y sur de la Argentina. Herpetofauna de las zonas áridas y semiáridas. Museo Regionale di Scienze Naturali, Torino, Monografie IV, 527 pp.
Cei, J.M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Museo Regionale di Scienze Naturali, Torino, Monografie, 14, 1–949.
Cei, J.M. & Scolaro, J.A. (1977) Herpetología patagónica. XIII. La identidad de Liolaemus goetschi y de la forma melanops del groupo Liolaemus fitzingerii, en Río Negro y Chubut. Physis, 36, 225–226.
Cei, J.M. & Scolaro, J.A. (1980) Two new subspecies of the Liolaemus fitzingerii complex from Argentina.Journal of Herpetology, 14, 37–43.
Cei, J.M. & Scolaro, J.A. (1983) Un nuevo arreglo taxonómico para los Liolaemus del grupo fitzingerii. Boletín de la Asociación Herpetológica Argentina, 1, 15–16.
Cei, J.M. & Scolaro, J.A. (1999) Speciation of the "darwinii complex" (genus Liolaemus, "patch group") in the southernmost area of its distribution (Reptilia: Tropiduridae). Revue francais de. Aquariologie, 26, 79–82.
Cei, J.M., Lescure, J. & Ortiz J.C. (1980) Rédecouverte de l’holotype de Proctotretus signifer Duméril and Bibron, 1837 (Reptilia, Iguanidae). Bulletin Museum National d’Historie Naturelle (Paris) Ser. 4, Sect. A (3), 919–925.
Díaz Gómez, J.M. (2007) Endemism on Liolaemus (Iguania: Liolaemidae) from the Argentine Puna. South American Journal of Herpetology, 2(1), 2007, 59–68.
Díaz Gómez, J.M. (2011) Estimating Ancestral Ranges: Testing Methods with a Clade of Neotropical Lizards (Iguania: Liolaemidae). Plos One 6(10), e26412.
Díaz Gómez, J.M. & Lobo, F. (2006) Historical Biogeography of a clade of Liolaemus (Iguania: Liolaemidae) based on ancestral areas and dispersal-vicariance analyisis (DIVA). Papéis Avulsos de Zoología, 46(24), 261–274
Duméril, A.M.C. & Bibron, G. (1837) Erpétologie générale ou histoire naturelle complète des reptiles. Ed. Roret, Paris 4, ii+571 pp.
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 herbivory. Proceedings of the National Academy of Sciences, 101, 16819–16824.
Etheridge, R.E. (1993) Lizards of the Liolaemus darwinii complex (Squamata: Iguania: Tropiduridae) in northern Argentina. Bollettino del Museo Regionale di Scienze Naturali Torino, 11, 137–199.
Etheridge, R.E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia:Squamata: Tropiduridae). American Museum Novitates, 3142, 1–34.
Etheridge, R.E. (2000) A review of the Liolaemus wiegmannii group Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293–352.
Etheridge, R. & Espinoza, R.E. (2000) Taxonomy of the Liolaeminae (Squamata: Iguania: Tropiduridae) and a semi-annotated bibliography. Smithsonian Herpetological Information Service, 126, 1–64.
Frost, D. (1992) Phylogenetic analysis and taxonomy of the Tropidurus group of lizards (Iguania: Tropiduridae). American Museum of Natural History Novitates, 3033, 68 pp.
Giribet, G. (2005) A review of: ‘‘TNT: Tree Analysis Using New Technology’’. Systematic Biology, 54, 176–178.
Goloboff, P. (1993) Estimating character weights during tree search. Cladistics, 9, 83–91.
Goloboff, P. Farris J. & Nixon, K. (2003) T.N.T.: Tree Analisis Using New Technology. Program and documentation, available from www.zmuc.dk/public/phylogeny
Goloboff, P.A. Farris, J.S. Källersjö, M. Oxelman, B. Ramírez, M. & Szumik, C.A. (2003a) Improvements to resampling measures of group support. Cladistics, 19, 324–332
Goloboff, P.A. Mattoni, C.I. & Quinteros, A.S. (2006) Continuous Characters Analyzed as Such. Cladistics, 22, 589–601.
Goloboff, P.A. Carpenter, J.M. Arias, J.S. & Miranda Esquivel, D.R. (2008) Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics, 24, 1–16.
Halloy, M. Etheridge, R. & Burghardt, G. M. (1998) To bury in the sand: phylogenetic relationships among lizard species of the boulengeri group, Liolaemus (Reptilia: Squamata: Tropiduridae), based on behavioral characters. Herpetological Monographs, 12, 1–37.
Halloy, M. Robles, C. & Cuezzo, F. (2006) Diet in two syntopic neotropical lizard species of Liolaemus (Liolaemidae) interspecifi c and intersexual diff erences. Revista Española de. Herpetología, 20, 47–56.
Halloy, M. Guerra, C. & Robles, C. (2007) Nuptial Coloration In Female Liolaemus quilmes (Iguania: Liolaemidae): Ambiguity And Keeping Males Interested? Bulletin of the Maryland Herpetological Society, 43 (3), 110–118.
Hovenkamp, P.H. (2004) Review of TNT. Cladistics, 20, 378–383.
Kacoliris F., Williams, J., Sánchez Véliz, G. & Rafael, A. (2008) Observaciones sobre el uso de cavidades en la arena por parte de la lagartija de los médanos (Liolaemus multimaculatus). Cuadernos de Herpetología, 22 (2), 87–89
Koslowsky, J. (1898) Enumeración sistemática y distribución de los reptiles argentinos. Revista Museo La Plata, 8, 161–200.
Kozykariski, M.L. Belver, L.C. & Ávila, L.J. (2011) Diet of the desert lizard Liolaemus pseudoanomalus (Iguania: Liolaemini) in northern La Rioja Province, Argentina. Journal of Arid Environments, 75, 1237–1239.
Labra, A., Carazo, P., Desfilis, E. & Font, E. (2007) Agonistic interactions in a Liolaemus lizard: structure of head bob displays. Herpetologica, 63 (1), 11–18.
Laurent, R.F. (1983) Contribución al conocimiento de la estructura taxonómica del género Liolaemus Wiegmann (Iguanidae). Boletín de la Asociación Herpetológica Argentina, 1, 15–18.
Laurent, R.F. (1985) Segunda contribución al conocimiento de la estructura taxonómica del género Liolaemus Wiegmann (Iguanidae). Cuadernos de Herpetología, 1, 1–37.
Lobo, F. (2001) A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae). Herpetological Journal, 11 (4), 137–150.
Lobo, F. (2005) Las relaciones filogenéticas dentro del grupo chiliensis (Iguania: Liolaemidae: Liolaemus): sumando nuevos caracteres y taxones. Acta Zoológica Lilloana, 49(1–2), 67–89.
Lobo, F. & Espinoza, R.E. (1999) Two new cryptic species of Liolaemus (Iguania: Tropiduridae) from northwestern Argentina: resolution of the purported reproductive bimodality of Liolaemus alticolor alticolor. Copeia, 1, 122–140.
Lobo, F., Espinoza, R. & Quinteros, S. (2010) A critical review and systematic discussion of recent classification proposals for liolaemid lizards. Zootaxa, 2549, 1–30.
Minoli, I. & Ávila, L.J. (2011) Liolaemus xanthoviridis (Rawson Lizard). Herpetological Review, 42(2), 2011
Morando, M. (2004) Sistemática y Filogenia de grupos de Especies de los géneros Phymaturus y Liolaemus (Squamata: Tropiduridae: Liolaemidae). Tesis Doctoral Universidad Nacional de Tucumán, 249 pp.
Moreno Azócar, L. & Acosta, J.C. (2011) Feeding Habits of Liolaemus cuyanus (Iguania: Liolaemidae) from the Monte Biogeographic Province of San Juan, Argentina. Journal of Herpetology, 45(3), 283–286
Nori J., Abdala, C.S. & Scrocchi, G.J. (2010) Liolaemus goetschi (Iguania: Liolaemidae): redescription and phylogenetic relationships within the L. boulengeri group. Zootaxa, 2440, 49-59.
Peters, J.A. & Donoso-Barros, R. (1970) Catalogue of the Neotropical Squamata: Part II. Lizards and amphisbaenians.Bulletin of the United States National Museum, 297, 1-293.
Quinteros, A.S. & Abdala, C.S. (2011) A new species of Liolaemus of the Liolaemus montanus section (Iguania: Liolaemidae) from Northwestern Argentina. Zootaxa, 2789, 35–48
Quinteros, A.S. Abdala, C.S. & Lobo, F.J. (2008) Redescription of Liolaemus dorbignyi Koslowsky, 1898 and description of a new species of Liolaemus (Iguania: Liolaemidae). Zootaxa, 1717, 51–67.
Schulte, J.A., Macey, J.R. Espinoza, R.E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring andean vicariance and dispersal. Biological Journal of the Linnean Society, 69, 75‑102.
Scolaro, J.A. & Cei, J.M. (1997) Systematic status and relationships of Liolaemus species of the archeforus and kingii groups: a morphological and taxonumerical approach (Reptilia: Tropiduridae). Boll. Mus. Reg. Sci. Nat. Torino, 15 (2), 369–406.
Scolaro, J.A. Cei, J.M & Arias de Reyna, L. (1985) La identidad de las especies del grupo Liolaemus fitzingerii-melanops por medio del análisis discriminante de caracteres morfológicos (Sauria: Iguanidae).Historia Natural (Corrientes, Argentina), 5, 13–22
Smith, H.M. (1946) Handbook of Lizards.Comstock Publ. Co., New York, xxi + 557 pp.
Tulli M.J., Abdala, V. & Cruz, F.B. (2011) Relationships among morphology, clinging performance and habitat use in Liolaemini lizards. Journal of Evolutionary Biology, 24 (4), 843-855
Valdecantos, M.S. & Lobo, F. (2007) Dimorfismo sexual en Liolaemus multicolor y L. irregularis (Iguania: Liolaemidae). Revista Española de Herpetología, 21, 55–69.
Valdecantos, S., Lobo, F. & Martinez, V. (2007) Estimación de edades, tamaño corporal y adquisición de la madurez sexual en dos especies de Liolaemus (Iguania: Liolaemidae). Cuadernos de Herpetología, 21,31–44.