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Species Limits in the Genus Gerrhonotus (Squamata: Anguidae)

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Species Limits in the Genus Gerrhonotus (Squamata: Anguidae) Species Limits in the Genus Gerrhonotus (Squamata: Anguidae) David A. Good Herpetological Monographs, Vol. 8. (1994), pp. 180-202. Stable URL: http://links.jstor.org/sici?sici=0733-1347%281994%298%3C180%3ASLITGG%3E2.0.CO%3B2-8 Herpetological Monographs is currently published by Herpetologists' League. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/herpetologists.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers, and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take advantage of advances in technology. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Wed Dec 12 18:51:01 2007 180 HERPETOLOGICAL MONOGRAPHS [No.8 duction by Uta stansburiana (Reptilia, Lacertilia, parative life histories of two syntopic sceloporine Iguanidae) in southern Nevada. J. Herpetol. 10: lizards. Copeia 1986:l-18. 123-128. TINKLE,D. W., AND L. N. IRWINE. 1965. Lizard MOCK,D., H. DRUMMOND,AND C. H. STINSON.1990. reproduction: Refractory period and response to Avian siblicide. Am. Sci. 78:438-449. warmth in Uta stansburiana females. Science 148: MOLL, E. 0. 1979. Reproductive cycles and ad- 1613-1614. aptations. Pp. 305-331. In M. Harless and H. Mor- TINKLE,D. W., H. M. WILBUR,AND S. G. TILLEY. lock (Eds.), Turtles: Perspectives and Research. John 1970. Evolutionary strategies in lizard reproduc- Wiley and Sons, New York. tion. Evolution 24:55-74. NUSSBAUM,R. A. 1981. Seasonal shifts in clutch size TRIVERS,R. L., AND D. E. WILLARD.1973. Natural and egg size in the side-blotched lizard, Uta stans- selection of parental ability to vary the sex ratio of buriana Baird and Girard. Oecologia 49:8-13. offspring. Science 179:90-92. SCHOENER,T. W. 1971. Theory of feeding strate- VINEGAR,M. B. 1975. Demography of the striped gies. Ann. Rev. Ecol. Syst. 2:369-404. plateau lizard, Sceloporus virgatus. Ecology 56:172- SEXTON,0. J., AND 0. TURNER.1971. The repro- 182. ductive cycle of a neotropical lizard. Ecology 52: VITT,L. J. 1978. Caloric content of lizard and snake 159-164. (Reptilia) eggs and bodies and the conversion of SMITH,C. S., AND S. D. FRETWELL.1974. The op- weight to caloric content. J. Herpetol. 12:65-72. timal balance between size and number of off- . 1981. Lizard reproduction: Habitat spec- spring. Am. Nat. 108:499-506. ificity and constraints on relative clutch mass. Am. STAMPS,J. A,, AND S. K. TANAKA.1981. The influ- Nat. 117:506-514. ence of food and water on growth rates in a tropical . 1990. The influence of foraging mode and lizard (Anolis aeneus). Ecology 62:33-40. phylogeny on seasonality of tropical lizard repro- STEARNS,S. C. 1984. The effects of size and phy- duction. Papkis Avulsos Zool. 37:107-123. logeny on patterns of covariation in the life history VITT,L. J., AND J. D. CONGDON.1978. Body shape, traits of lizards and snakes. Am. Nat. 123:56-72. reproductive effort, and relative clutch mass in liz- TINKLE,D. W. 1967. The life and demography of ards: Resolution of a paradox. Am. Nat. 112:595- the side-blotched lizard, Uta stansburiana. Misc. 608. Publ. Mus. Zool. Univ. Michigan 132:l-182. VITT, L. J., AND H. J. PRICE. 1982. Ecological and TINKLE,D. W., AND R. E. RALLINGER.1972. See- evolutionary determinants of relative clutch mass loporus undulatus: A study of the intraspecific in lizards. Herpetologica 38:237-255. comparative demography of a lizard. Ecology 53: 570-584. Accepted: 11 March 1993 Associate Editor: Stanley Hillman TINKLE,D. W., AND A. E. DUNHAM.1986. Com- Herpetological Monographs, 8, 1994, 180-202 O 1994 by The Herpetologists' League, Inc. SPECIES LIMITS IN THE GENUS GERRHONOTUS (SQUAMATA: ANGUIDAE) DAVIDA. GOOD Museum of Natural Science, Louisiana State University, Baton Rouge, LA 70803, USA ABSTRACT:Gerrhonotus liocephalus is demonstrated to consist of at least three species rather than the single polytypic species currently recognized. Of the seven currently recognized subspecies, G. 1. aguayoi, G. 1, infernalis and G. 1. taylori become G. infernalis; G. 1. ophiurus and G. 1, loweryi become G.ophiurus; and G.1, austrinus and G.1. liocephalus retain the name G.liocephalus. No subspecies are recognized within any of these three species. Certain populations from western Mexico (Durango, Sinaloa, Jalisco, and Colima) remain of uncertain identity, but are tentatively referred to as G. cf, liocephalus. Evidence for these conclusions is derived from an analysis of scalation, coloration, and morphometric variation. The taxonomic history in Gerrhonotus also is discussed Key words: Alligator lizard; Anguidae; Gerrhonotus;Mexico; Species; Systematics; Texas THEgenus Gerrhonotus at present con- other hand, is among the most widely dis- tains two recognized species. Gerrhonotus tributed alligator lizards (Anguidae: Ger- lugoi is a poorly known form restricted to rhonontinae). Geographic variation in the the vicinity of Cuatro Ciknegas, Coahuila, latter taxon has led to the description of Mexico. Gerrhonotus liocephalus, on the seven subspecies: aguayoi, austrinus, in- 19941 HERPETOLOGICAL MONOGRAPHS 181 fernalis, liocephalus, loweryi, ophiurus, apparently were unaware of the descrip- and taylori, with another suggested but not tion of G. liocephalus by Wiegmann and formally described (Smith, 1984). Several did not compare the two taxa. Wiegmann workers have discussed differences in sca- (1834), however, recognized that his G. lation, body proportions, and color pattern liocephalus was identical to S. ventralis. among these forms, but since 1900 this Because he decided that the name "lio- discussion has been restricted to the con- cephalus" ("smooth head") was appropri- text of a single polytypic species. In this ate for the juvenile he had originally de- paper, I will discuss the history of intra- scribed but not for adult specimens such specific taxonomy in the broadly defined as those examined by Peale and Green, G. liocephalus in order to provide a back- Wiegmann (1834) changed the name of ground for the currently accepted specific the species to G, tessellatus. This latter and subspecific limits. I will then re-ana- name was in common use (e.g.,Cope, 1866, lyze the patterns of variation in scalation, 1878; Dumkril and Bibron, 1839; Dumkril morphometrics, and color pattern within and Dumbril, 1851; Gray, 1838, 1845; and among the currently recognized sub- O'Shaughnessy, 1873) until Bocourt (1878) species, and provide evidence for the di- resurrected liocephalus. vision of G. liocephalus into multiple spe- The first Texan specimen, from the can- cies. yon of the Devil's River, was described by In discussing the results of this analysis, Baird (1858) as G. infernalis. Baird did I will use the first six subspecies names not compare his new species with either listed above for ease of comparison with G. liocephalus/G. tessellatus or Scincus previous papers, although I will conclude ventralis. later in the paper that some of them should Cope (1866)described G. ophiurus from be synonymized. The description of Orizaba, Veracruz, as differing from G. aguayoi (Contreras Arquieta, 1989) ap- tessellatus by having a longer tail, shorter peared only in an abstract from a scientific limbs, and a different arrangement of the meeting and hence is of questionable va- head scales. Cope (p. 321) summarized lidity. Because no type specimen was des- what was then known about variation in ignated, the populations referable to it are Gerrhonotus as follows (with terminology unclear and I cannot include it in the char- modified to match that used in the present acter analysis below. I will refer to three paper): unnamed western groups of populations as "western isolates" in presenting the results One preocular; canthal/loreal series consist- and then hypothesize possible species ing of an anterior canthal, an anterior lo- placement in the Discussion. real, and a posterior cantholoreal; ad- pressed limbs separated by the length of Intraspeciflc Taxonomy in Gerrhonotus the hind leg; venter immaculate; tail rel- liocephalus (sensu lato) atively short . G. ventralis No preoculars; canthal/loreal series consist- In 1828, A. F. A. Wiegmann described ing of an anterior canthal, an anterior lo- six species of gerrhonotine lizards on the real, and a posterior cantholoreal; ad- basis of a set of specimens collected by F. pressed limbs separated by length of the Deppe and donated to the Berlin Museum. forearm; dorsum brown with ten cross- Among the specimens described by Wieg- bands; venter black-spotted; .. .. mann was a single juvenile he named Ger- . .. G. tessellatus [=G. liocephalus] rhonotus liocephalus. Wiegmann provid- Two preoculars; three loreals present, no ed no type locality, although the holotype cantholoreal; anterior internasal contact- was clearly from Mexico (as suggested by ing the anterior supralabial; adpressed limbs separated by humerus length; dor- its inclusion by Wiegmann, 1834, in his sum red, with ten light V-shaped cross- Herpetologia Mexicana). bands; venter black-spotted; tail length Soon after the appearance of Wieg- 2.75 times snout-vent length .
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