NovitatesAMERICANt MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3207, 60 pp., 27 figures, 9 tables October 23, 1997 The Blue and Green Whiptail Lizards (Squamata: Teiidae: Cnemidophorus) of the Peninsula de Paraguana, Venezuela: Systematics, Ecology, Descriptions of Two New Taxa, and Relationships to Whiptails of the Guianas ALLAN L. MARKEZICH,1 CHARLES J. COLE,2 AND HERBERT C. DESSAUER3 CONTENTS Abstract ...................................................................... 2 Resumen ...................................................................... 3 Introduction .................................................................... 4 Acknowledgments ............................................................... 5 Methods ......... .............................................. 6 Results and Discussion ......................... ................................. 11 Evidence from Karyotypes ................. .................................... 11 Evidence from Biochemical Genetics ........................................... 11 Are the Green and the Blue Cnemidophorus from the Paraguana Peninsula Two Different Species9. ........................................................ 13 'Field Associate, Department of Herpetology, American Museum of Natural History; and Professor, Department of Natural Sciences and Engineering, Black Hawk College, Moline, IL. 2Curator, Department of Herpetology, American Museum of Natural History. 3Research Associate, Department of Herpetology, American Museum of Natural History; and Professor Emeritus, Department of Biochemistry and Molecular Biology, Louisiana State University Medical Center, New Orleans, LA. Copyright © American Museum of Natural History 1997 ISSN 0003-0082 / Price $7.20 2 AMERICAN MUSEUM NOVITATES NO. 3207 Is One of the Species from the Paraguana Peninsula a Local Population of C. lemniscatus9.13Iemisctus ............................................ Is One of the Species from the Paraguana Peninsula a Local Population of C. gramivagus9.13gramiagus ............................................ Is One of the Species from the Paraguana Peninsula The Second Ancestor of the Unisexual C. cryptus9 ......................... 15 Evidence from Morphology: Comparisons Among the Blue and the Green Lizards and C. lemniscatus from the Guianas .......... ................... 17 Sexual Dimorphism ............................. 17 Univariate Comparisons ............................. 17 Multivariate Analysis ............................. 19 Variation in Color Pattern ............................. 21 Ecology ..................................................................... 25 Diet ....................................................................... 36 Reproduction ............................................................... 37 Taxonomic and Evolutionary Considerations ..................................... 38 Descriptions of the New Taxa .................................................. 42 Rapid Identification of Specimens .............................................. 50 Additional Names and Relevant Type Specimens ................................. 50 The Types of C. scutata ..................................................... 51 The Lectotype of C. lemniscatus ............................................. 51 Summary and Conclusions ..................................................... 53 References .................................................................. 55 Appendix-Specimens Examined ................................................ 59 ABSTRACT Two color morphs of whiptail lizards (Cne- cies is compared with populations of C. lemnis- midophorus), one brilliant blue and one bright catus from northern Venezuela and northeastern green, occur in distinctly different habitats on Colombia and with Cnemidophorus 1. gaigei of the Paraguana Peninsula, Venezuela. Multidis- Colombia, which is currently recognized as a ciplinary analyses (karyotypes, protein electro- synonym of Cnemidophorus 1. lemniscatus. In phoresis, color pattern, scalation, using univari- addition, we discuss nomina dubia in the syn- ate and multivariate statistical techniques) re- onymy of C. lemniscatus, we describe and il- vealed that the blue and the green lizards rep- lustrate the types of C. scutata (a synonym of resent two distinctively different diploid C. 1. lemniscatus), we illustrate the lectotype of bisexual species. The green lizard is an unde- C. lemniscatus, and we present a table of taxo- scribed species of Cnemidophorus, which also nomic characters for identifying the valid taxa occurs in other areas of Falcon, Venezuela and that are described and reviewed. on the Guajira Peninsula, Colombia, in open The relationships of both new taxa to the geo- dune or desert scrub communities. Comparisons graphically nearest congener, Cnemidophorus with known C. lemniscatus from the Guianan arubensis of Aruba Island, which is only 30 km Region suggest that the blue lizard, which oc- north of the Paraguana Peninsula, are discussed, curs in less open habitats such as tropical thorn but they remain obscure. A small population of woodland, is best treated as an undescribed sub- lizards considered by previous workers to be species of C. lemniscatus. Although its occur- Cnemidophorus lemniscatus in sympatry with rence off the Paraguana Peninsula is suspected C. arubensis at one locality on Aruba Island also, this remains to be documented. probably is the green species described here as Both new taxa are named and diagnosed. Col- new. or photographs and details of morphological The two species of Cnemidophorus on the and genetic variation for the new taxa are pre- Paraguana Peninsula are mainly allopatric or sented and correlated with data from C. lemnis- narrowly parapatric in their microgeographic catus of the Guianan Region. The new subspe- distribution but sympatric at a few localities 1997 MARKEZICH ET AL.: BLUE AND GREEN WHIPTAIL LIZARDS OF PARAGUANA 3 with disturbed habitats. They do not appear to preferences and diet (the green taxon is primar- interbreed and are allotopic in areas of sympa- ily herbivorous), and they are similar in repro- try. These taxa differ from each other in habitat ductive biology. RESUMEN Para muchas areas del neotr6pico falta informa- como sinonimo de Cnemidophorus 1. lemniscatus. ci6n sobre la taxonomia, ecologia e historia natural Ademas, lo tratamos nomina dubia en la sinoni- de los lagartos indigenas, la cual es fundamental mia; describimos los tipos de C. scutata (un si- para poder entender la historia de la diferenciaci6n nonimo de C. 1. lemniscatus), ilustramos el lec- y el significado de las distribuciones actuales y su totipo de C. lemniscatus y presentamos una tabla diversidad. La arida regi6n de tierra baja del no- de caracteres taxon6micos para identificar los taxa roeste de Venezuela es uno de esos tipos de zonas, va'lidos, descritos y revisados. de modo que uno de nosotros (ALM) comenz6 el La relaci6n de ambos taxa nuevos con el con- estudio de la ecologia y diversidad de la herpeto- genere mas cercano geograficamente, el Cnemi- fauna de la peninsula de Paraguana en 1989. dophorus arubensis de la isla de Aruba, que esta Las observaciones de los lagartos (Cnemidopho- situada solamente a 30 km al norte de la peninsula rus) de alli, revelo una relacion sorprendente entre de Paraguana, es poco clara. Se precisaran datos el colorido y la ecologia. Se daban dos morfos dis- cariotfpicos y otros datos geneticos de este ultimo tintos de color en habitats diferentes. Un morfo in- para poder resolverlo. El aislamiento de C. aru- cluia lagartos de un azul brillante, el otro, lagartos bensis se trata en relaci6n con la paleograffa de de un verde brillante. Sin embargo, se pensaba que Aruba y de la penfnsula de Paraguand durante el solamente una de las especies, el Cnemidophorus ultimo maximum glaciar. Una pequefia poblaci6n lemniscatus se daba en esta area de Venezuela. Estas de lagartos que investigadores anteriores consi- especies y los congeneres de la regi6n de Guyana deraron que fuera Cnemidophorus lemniscatus en (al norte del Rfo Amazonas y al este del Orinoco y simpatria con C. arubensis en una localidad de la del Rio Negro) descritos recientemente, han sido isla de Aruba, es probablemente la especie verde considerados mas bien generales en lo que al habitat que se describe aqui como nueva. se refiere. El Cnemidophorus verde de la peninsula de Par- Anailisis multidisciplinarios (cariotipos, electro- aguand se da en habitats abiertos con vegetacion foresis de proteinas, colorido y escamacion, usan- baja, tal como dunas de arena o matorrales de do tecnicas estadisticas univariadas y multivaria- rastrojo. Es un herbivoro facultativo como otros das) revelaron que los lagartos azules y lagartos congeneres proximos de la peninsula, una adap- verdes de la peninsula de Paraguani son dos es- tacion que quiza's se desarroll6 como respuesta al pecies diploides y bisexuales claramente distintas. severo medioambiente con reservas de comida El lagarto verde es una especie de Cnemidophorus impredecibles. El Cnemidophorus azul se asocia no descrita, que se da en otras areas de Falc6n, con haibitats mas sombreados con vegetaci6n mas en Venezuela y en la peninsula de Guajira, en Co- alta y espesa, tal como el matorral espinoso trop- lombia. Las comparaciones con la conocida C. ical o comunidades de bosques tropicales muy se- lemniscatus de la regi6n de Guyana sugieren que cos, y es carnivoro. Es posible que su distribuci6n