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Novitatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

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Novitatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y NovitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3081, 30 pp., 11 figures, 2 tables December 27, 1993 Unisexual and Bisexual Whiptail Lizards of the Cnemidophorus lemniscatus Complex (Squamata: Teiidae) of the Guiana Region, South America, with Descriptions of New Species CHARLES J. COLE1 AND HERBERT C. DESSAUER2 CONTENTS Abstract ..................... 2 Introduction ..... 2 Acknowledgments ..................... 3 Materials and Methods ..................... 4 Results and Discussion ..................... 5 Karyotypes .......................5 Biochemical Genetics ..................... 5 Hybrid Origins ..................... 10 Reproduction ..................... 11 Evolutionary Scenano ..................... 12 Implications for Biodiversity ..................... 13 Taxonomy ..................... 15 Alternative Treatments ..................... 25 Summary and Conclusions ..................... 26 References ..................... 27 Appendix-Specimens Examined ................................................ 29 ' Curator, Department of Herpetology and Ichthyology, American Museum of Natural History. 2 Research Associate, Department ofHerpetology and Ichthyology, American Museum ofNatural History; Professor Emeritus, Department of Biochemistry and Molecular Biology, Louisiana State University Medical Center, New Orleans, LA. Copyright © American Museum of Natural History 1993 ISSN 0003-0082 / Price $3.25 2 AMERICAN MUSEUM NOVITATES NO. 3081 ABSTRACT At least four species have been masquerading A revised diagnosis is presented for C. lemnis- under the name Cnemidophorus lemniscatus in catus, a male lectotype is designated, and the type northern South America. These species were re- locality is restricted to an area in Suriname with vealed clearly by karyotypic and electrophoretic a bisexual population oflizards similar to the Lin- analyses of samples from unisexual and from bi- naean syntypes and consistent with their suspected sexual populations. The four taxa include two dip- provenance. The second bisexual species appears loid bisexual species and two unisexual cryptic to be C. gramivagus McCrystal and Dixon, for species (one diploid and one triploid). The uni- which a revised diagnosis is presented also. New sexual species are parthenogenetic clones that ul- diagnoses and names are presented for two uni- timately originated from hybrids among the bi- sexual species, a diploid based on specimens from sexual species (the triploids stemming from Venezuela (with paratypes from Brazil also), and backcross hybridization). There are several differ- a triploid based on specimens from Surname. Al- ent diagnosable diploid clones, at least two ofwhich ternative taxonomic treatments for the various probably had origins from separate hybridization diploid unisexual clones found in Brazil are dis- events at different localities. Several other diag- cussed, but none of these is named as a separate nosable diploid clones probably stemmed from species here. For now, we treat all of the diploid mutations within one or more established parthe- clones as one species. nogenetic lineages of very similar hybrid origin. INTRODUCTION Cnemidophorus lemniscatus (Linnaeus) is catus from Brazil by comparing products of a widespread and generally abundant and 57 presumed structural gene loci in types B conspicuous whiptail lizard in much of Cen- through E, using protein electrophoresis. The tral America and South America, mostly in hypothesis was strongly supported, as bisex- bisexual populations (gonochoristic, with ual types D and E had relatively low hetero- separate sexes; fertilization is required for re- zygosity indices and fixed or nearly fixed dif- production). Vanzolini (1970) first reported ferences from each other at 11 of the loci that some populations along the Amazon analyzed. In contrast, the unisexual types B River in Brazil are bisexual and others are and C had relatively high heterozygosity and unisexual (all-female). genotypes consistent with their having a hap- Chromosome studies by Peccinini-Seale loid set of genes each from bisexual types D and Frota-Pessoa (1974) revealed five slightly and E, with minor exceptions. Sites et al. different karyotypes among the Brazilian C. (1990) proposed that at least three species are lemniscatus, which they designated alpha- involved in this complex along the Amazon betically, A through E. Types A-C are minor River in Brazil: bisexual type D, bisexual type variants characterizing separate unisexual E, and the diploid unisexuals ofhybrid origin. populations, all diploids, differing from each A companion study ofmitochondrial DNA other primarily in a centromere position (Vyas et al., 1990) showed that the Brazilian among the second largest pair of macrochro- unisexuals have only one type ofmtDNA and mosomes and in number of microchromo- it is most similar to that occurring in certain somes. Types D and E characterize the bi- bisexual type D lizards (i.e., type D was the sexual populations, which differ from each maternal ancestor). There is variation in other only in centromere position on chro- mtDNA in type D, and none of the type D mosome number 1 (largest pair). Type C of samples examined matched perfectly the the unisexuals has a karyotype that could be mtDNAs of the unisexuals. Nevertheless, composed of one haploid complement each none of the unisexuals had mtDNA similar from bisexual types D and E; i.e., it could to that ofbisexual type E (i.e., type E was the have had a hybrid origin. Unisexual types A paternal ancestor). and B could have been derived from unisex- Separate bisexual and unisexual popula- ual type C by chromosomal aberrations. tions of C. lemniscatus also occur in Suri- Sites et al. (1990) tested the hybrid origin name (Hoogmoed, 1973), but these unisex- hypothesis for diploid unisexual C. emnis- uals are triploids rather than diploids (Serena, 1993 COLE AND DESSAUER: UNISEXUAL AND BISEXUAL WHIPTAIL LIZARDS 3 1985; Dessauer and Cole, 1989). Studies of type D bisexual C. lemniscatus from Suri- karyotypes and protein electrophoresis (46 name and Venezuela compare favorably with loci tested) suggested that the bisexual dip- those from Brazil, as indicated by karyotypes loids in Suriname are similar to type D of and protein electrophoresis? (2) Do the uni- Brazil (same taxon), and the triploids had a sexual samples from Brazil, Suriname, and hybrid origin involving two genomes of type Venezuela have basically the same genotypes, D and one of type E (Dessauer and Cole, including the alleles at heterozygous loci that 1989). This probably resulted from back- were inherited from bisexual type E? (3) What crossing ofa diploid unisexual oftype C (orig- is the minimal number ofspecies represented inally of type D x type E origin) with a male by the samples compared here, and what of type D; i.e., a double hybrid origin of [D names should be applied to them? A review x E] x D. of these questions improves our knowledge Electrophoretic analyses of the samples ofbiodiversity and its origins in Neotropica. from Suriname and from Brazil were con- ducted separately in different laboratories ACKNOWLEDGMENTS (Dessauer and Cole, 1989; Sites et al., 1990, Permits and considerable assistance with respectively), so cross-correlation gels com- logistics in the field were provided by S. A. paring all of the samples together did not J. Malone, F. L. J. Baal, and M. H. S. Miskin exist. Evidence that the genomes of type D (all of the Suriname Forest Service), Giu- and E were basically the same in unisexual seppe Colonnello (Museo de Historia Natural lizards from Brazil and Suriname was pro- La Salle, Caracas, Venezuela), and Karen Pil- vided only by the karyotypes (Dessauer and grim (Wildlife Services Division, Depart- Cole, 1989: fig. 7C), which do not resolve ment of Agriculture, Guyana). We were as- genetic data as well as protein electrophoresis sisted in the field in Suriname by Jeffrey Cole, does. To determine whether the same alleles in Venezuela by Carlos Guaveco and Maria at structural gene loci are shared by lizards Jose Praderio (Museo de Historia Natural La in Brazil and Suriname (type D bisexuals and Salle), and in Guyana by Bruce Hoffman. diploid and triploid unisexuals), samples Fieldwork in Guyana in 1992 was con- would have to be subjected to protein elec- ducted through the Biological Diversity of trophoresis on the same gels. the Guianas Program (BDGP), coordinated While the two independent electrophoretic in Guyana by Deputy Vice-Chancellor Mal- reports discussed above on lizards from Su- colm Rodrigues and Dean Indarjit Ramdass riname and Brazil were in press, CJC, Carlos ofthe University ofGuyana (UG), Turkeyen Guaveco, Maria Jose Praderio, and Carol R. Campus, Georgetown. Mike Tamessar ofUG Townsend collected two new samples of the also greatly facilitated this work. Important C. lemniscatus complex in Venezuela. Spec- logistical support was provided by personnel imens from San Ignacio de Yuruani (State of serving the BDGP, including Vicki Funk, Bolivar) included both sexes, whereas those Bruce Hoffman, Carol Kelloff, and Moham- from Icabaru (State ofBolivar) included only med Ameer (the females, although no unisexual populations "Ambassador of Bourda"). were known for Venezuela. Additional logistical support was provided For the present report, we karyotyped the by Malcolm Chan-A-Sue (Torong Guyana), new samples from Venezuela and we ana- Walter Lachman (National Dairy Develop- lyzed together
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