Richard Etheridge

Two New Species of the Lizard Genus Liolaemus (Squamata: Liolaemidae) from Northern Patagonia, with Comments on Liolaemus rothi

Abstract A number of undescribed species of the iguanian lizard genus Liolaemus, formerly referred to Liolaemus rothi and Liolaemus boulengeri, have been identified in the northern Patagonian region of Argentina. Descriptions of two of the species are presented, along with their distributions and natural history. Liolaemus rothi is redescribed, a lectotype is designated, its likely provenience is discussed, and its distribution and natural history are given.

PHYLOGENETIC RELATIONSHIPS OF HOPLOCERCID LIZARDS: CODING AND COMBINING MERISTIC, MORPHOMETRIC, AND POLYMORPHIC DATA USING STEP MATRICES

Hoplocercidae is a small (10 species, 3 genera), poorly known but pivotal family of Central and South American iguanian lizards that has never been the subject of a focused phylogenetic study or systematic revision. We undertake the first rigorous phylogenetic analysis of hoplocercid lizards. We also use our analysis to demonstrate how meristic, morphometric, and polymorphic morphological characters can be coded and combined for phylogenetic analyses using step matrices, which allow continuous variation to be treated as continuous. Parsimony analysis of 46 informative external and skeletal characters (17 qualitative and fixed, 19 qualitative and polymorphic, 8 meristic, and 2 morphometric) yields very different topologies, depending on how the meristic characters are scaled (weighted). Use of between-state scaling yields a phylogeny in which Hoplocercus is at the base of the hoplocercid tree, and Morunasaurus is paraphyletic with respect to a monophyletic Enyalioides. Scaling between characters produces a tree in which Enyalioides is paraphyletic with respect to a clade containing Morunasaurus and Hoplocercus, and Morunasaurus is paraphyletic with respect to Hoplocercus. We also propose a third, “mixed” approach to scaling, which we marginally prefer over the other two methods. This method yields a tree in which Hoplocercus and a monophyletic Morunasaurus make up the sister group to a monophyletic Enyalioides. We discuss the implications of these results for hoplocercid biogeography and evolutionary ecology, tropical speciation, and the phylogenetic analysis of morphological data.

Nueva especie de lagartija del norte de Chile, Liolaemus molinai (Reptilia: Liolaeminae)

Se describe una nueva especie de lagartija del norte de Chile, perteneciente al genero Liolaemus y adscrita al grupo montanus. Los ejemplares de la nueva especie fueron capturados en febrero de 1999 en los Farellones de Tara, Provincia del Loa, Region de Antofagasta. Esta difiere de las especies del subgrupo boulengeri en que carece de un parche de escamas alargadas en la parte femoral de la extremidad posterior. Difiere de las demas especies del grupo montanus por presentar un gran numero de escamas pequenas, lisas y redondeadas en la zona dorsal del cuerpo, y escamas cefalicas distintivas. De las especies con patron de escamacion similar, la nueva especie puede ser diferenciada por su moderado tamano, cola mas corta que la longitud cabeza-cloaca y patron de coloracion distintivo. Es de habitos psammofilos y se encuentra sobre los 4.000 m de altura. Se amplificaron y secuenciaron 1.775 pb de los genes mitocondriales ND1, ND2 y COI del holotipo, alotipo y un paratipo de la nueva especie. Fueron comparadas con dieciseis especies del genero Liolaemus, cuatro de ellas del grupo chiliensis y doce del grupo montanus. Ademas fue usado como grupo externo un ejemplar del genero Phymaturus. Las secuencias fueron alineadas y analizadas en los programas Clustal W, Mac Clade, PAUP y MEGA2. El numero de sitios variables filogeneticamente informativos fue de 453. La divergencia genetica de la nueva especie con las congenericas del grupo chiliensis fue entre 0,1780 y 0,1985 unidades K2P, mientras que con las especies del grupo montanus fue de 0,0676 y 0,0857 unidades K2P. Con respecto al analisis de Maxima Parsimonia, se detecto el arbol mas corto con 1.682 pasos y con un Indice de Consistencia de 0,68, con una topologia basicamente similar a la obtenida en Maxima Verosimilitud y Distancia. El analisis de componentes principales permite diferenciar a la nueva especie de otras seis del grupo montanus, principalmente por las medidas corporales, las que explican mas del 84 % de la varianza. Tanto el analisis morfologico como los datos moleculares muestran claramente que esta es una nueva especie

The Type Locality of Phymaturus palluma (Molina, 1782) (Reptilia: Iguania: Liolaemidae) and the Status of Phymaturus adrianae Pereyra, 1992 and Other Unnamed Populations

Abstract. With the recent designation of the holotype of Centrura flagellifer Bell as theneotype of Phymaturus palluma Molina by the International Commission on Zoological Nomenclature, the type locality of P. palluma became problematic. It is likely that Charles Darwin collected the holotype of C. flagellifer during his round trip journey between Santiago, Chile and Mendoza, Argentina. A detailed account of his journey is presented, as well as the conclusion that the type locality is Cordón del Portillo, Mendoza province, Argentina, where Darwin collected and briefly characterized a viviparous lizard. Here we provide a re-description of Darwins specimen and two morphological characters that distinguish it from other populations in Chile and Argentina previously assigned to this species. In November 2009, specimens from the locality where Darwin collected his “viviparous lizard” were collected, confirming the identity of the type. This study also reveals that P. gynechlomus should be considered a junior synonym of P. palluma. Populations of Phymaturus of the palluma group farther to the north, from the Sierra de Uspallata and southern San Juan province, are determined to be different lineages and should be described formally. The name P. adrianae has been applied to the species that occurs in the Sierra de Uspallata, but we show here that it has not been formally named and is therefore a nomen nudum. Chilean populations previously considered to be P. palluma should be described formally.

The Type Locality ofPhymaturus palluma(Molina, 1782) (Reptilia: Iguania: Liolaemidae) and the Status ofPhymaturus adrianaePereyra, 1992 and Other Unnamed Populations

Abstract. With the recent designation of the holotype of Centrura flagellifer Bell as theneotype of Phymaturus palluma Molina by the International Commission on Zoological Nomenclature, the type locality of P. palluma became problematic. It is likely that Charles Darwin collected the holotype of C. flagellifer during his round trip journey between Santiago, Chile and Mendoza, Argentina. A detailed account of his journey is presented, as well as the conclusion that the type locality is Cordón del Portillo, Mendoza province, Argentina, where Darwin collected and briefly characterized a viviparous lizard. Here we provide a re-description of Darwins specimen and two morphological characters that distinguish it from other populations in Chile and Argentina previously assigned to this species. In November 2009, specimens from the locality where Darwin collected his “viviparous lizard” were collected, confirming the identity of the type. This study also reveals that P. gynechlomus should be considered a junior synonym of P. palluma. Populations of Phymaturus of the palluma group farther to the north, from the Sierra de Uspallata and southern San Juan province, are determined to be different lineages and should be described formally. The name P. adrianae has been applied to the species that occurs in the Sierra de Uspallata, but we show here that it has not been formally named and is therefore a nomen nudum. Chilean populations previously considered to be P. palluma should be described formally.

Taxonomic History of the Iguanian Lizard Liolaemus pictus major Boulenger, with a Revalidation of Liolaemus capillitas Hulse

Abstract Núñez (2004) examined the syntypes of Liolaemus pictus major Boulenger in the British Museum of Natural History and concluded, without supporting data, that this taxon is a senior synonym of Liolaemus capillitas Hulse. We show that the evidence does not support Núñezs (2004) proposal. We first document the complex taxonomic history of L. p. major, the lack of a precise or even definitive type locality, and the implications of the latter on subsequent checklists and research. Second, we note differences between Boulengers (1885) type description of L. p. major and Hulses (1979) type description of L. capillitas. Third, we show that the syntypes of L. p. major photographed by Núñez (2004) do not exhibit the character states of L. capillitas. We conclude that L. capillitas is not a synonym of L. p. major. Based on the available evidence, the syntypes of L. p. major are probably assignable to Liolaemus elongatus from populations occurring in Patagonia (southern Argentina or adjacent Chile). However, the precise identity of the syntypes requires additional study and perhaps a type locality restriction or redefinition of L. elongatus, which is itself in a state of flux.