Regular issues
Special Issues

Southeastern Naturalist
    SENA Home
    Range and Scope
    Board of Editors
    Editorial Workflow
    Publication Charges

Other EH Journals
    Northeastern Naturalist
    Caribbean Naturalist
    Urban Naturalist
    Eastern Paleontologist
    Eastern Biologist
    Journal of the North Atlantic

EH Natural History Home

Identity, Reproduction, Variation, Ecology, and Geographic Origin of a Florida Adventive: Cnemidophorus lemniscatus (Rainbow Whiptail Lizard, Sauria: Teiidae)
Brian P. Butterfield, J. Brian Hauge, Anthony Flanagan, and James M. Walker

Southeastern Naturalist, Volume 8, Number 1 (2009): 45–54

Full-text pdf (Accessible only to subscribers.To subscribe click here.)


Site by Bennett Web & Design Co.
2009 SOUTHEASTERN NATURALIST 8(1):45–54 Identity, Reproduction, Variation, Ecology, and Geographic Origin of a Florida Adventive: Cnemidophorus lemniscatus (Rainbow Whiptail Lizard, Sauria: Teiidae) Brian P. Butterfield1, J. Brian Hauge2, Anthony Flanagan3, and James M. Walker4,* Abstract - We answer a longstanding question pertaining to Florida’s exotic herpetofauna— are the colorful lizards (Sauria: Teiidae, Cnemidophorus lemniscatus [Rainbow Whiptail]), introduced to Florida from the pet trade, representative of one or a complex of species? We performed detailed analyses of color pattern, scutellation, and meristic characters using 16 specimens from Miami-Dade County and compared these data to those based on samples of all members in the widely distributed neotropical Cnemidophorus lemniscatus complex comprising both parthenogenetic (C. cryptus and C. pseudolemniscatus) and gonochoristic (C. arenivagus, C. gramivagus, C. lemniscatus lemniscatus, C. lemniscatus splendidus, and C. nigricolor) species of lizards. We concluded that only the most widely distributed taxon in the complex, C. lemniscatus lemniscatus, is established in Miami-Dade County. A population of this form has existed in an area of the City of Miami for several decades near and in a human-created domain of buildings, houses, swales, railroad right-of-way, and untended areas with litter/ debris, introduced plants, and variable substrate mixtures of soil/pebbles/concrete/ asphalt. Based on comparisons of color pattern and morphological characters, the founders of the population of C. lemniscatus in Florida were most likely imported from Venezuela, Colombia, or an island in the Caribbean Sea. Introduction The population of colorful lizards (Sauria: Teiidae) introduced to Florida, and uncertainly identified by various authors as either one or a complex of species (Bartlett and Bartlett 1999, Conant and Collins 1998, Meshaka et al. 2004, Punzo 2001, Wilson and Porras 1983), in fact, belongs to the gonochoristic species Cnemidophorus lemniscatus (Linnaeus) (Rainbow Whiptail). The extensive neotropical range of this taxon includes parts of northern South America, Central America, and numerous islands in the Caribbean Sea (Cole and Dessauer 1993, Markezich et al. 1997, Montgomery et al. 2007, Peters and Donoso-Barros 1970). Rainbow Whiptail in Florida, which has become established in Miami-Dade County over 1200 km northeast of the nearest natural populations of the species on Islas de Utila and Roatán, Honduras, is inferred to represent descendants of individuals imported for the pet trade (Meshaka et al. 2004). 1Department of Biology, Freed-Hardeman University, Henderson, TN 38340. 2Department of Biology, Peninsula College, Port Angeles, WA 98362. 32900 North 72th Avenue, Hollywood, FL 33024. 4Department of Biological Sciences, University of Arkansas, Fayetteville, AR 72701. *Corresponding author - 46 Southeastern Naturalist Vol. 8, No. 1 Cnemidophorus lemniscatus may have been introduced to Florida in 1964 with release of 40 individuals in Hialeah, Miami-Dade County (King and Krakauer 1966). This species is still extant in Hialeah, despite a report to the contrary (Wilson and Porras 1983). An image of an adult male captured there in 1988 was recently posted online by the Center for North American Herpetology at (University of Kansas Natural History Museum [KU 209859]), B.P. Butterfield and J.B. Hauge collected eight specimens in the city in 1999, and A. Flanagan observed and photographed juveniles and adults in 2008 in Hialeah. In 1996 and 1999, B.P. Butterfield and J.B. Hauge also collected a total of four specimens of C. lemniscatus in Virginia Gardens, Miami-Dade County, during a general survey of exotic amphibians and reptiles in Florida. The specimens of this species used by Meshaka et al. (2004) in the B.P. Butterfield and Auburn University Museum (AUM) collections, four individuals listed herein from the Florida Museum of Natural History (FLMNH), and 30 lizards studied by Punzo (2001) for diet composition verified that some of the Miami-Dade County lizards are gonochoristic (i.e., sexual) rather than parthenogenetic (i.e., asexual). Nevertheless, presence of males in those samples did not preclude the possibility that some of the females belonged to a parthenogenetic species in the C. lemniscatus complex. One of us (J.M. Walker), assessed color pattern, scutellation, and meristic variation for 16 lizards from Florida and compared these data to those for each of the other members of the C. lemniscatus complex. Herein, we use relevant parts of those comparisons to demonstrate that only gonochoristic C. lemniscatus is represented among the Floridian lizards examined in this study. We also report data on body size, many aspects of scutellation, reproduction, and ecology, and comment on the origin of founders, of this Florida adventive. Methods We measured the snout–vent length (SVL) of specimens of C. lemniscatus to the nearest mm (Table 1). Terminology pertaining to ontogenetic changes in dorsal color pattern in the Florida population includes: four pairs of palecolored primary stripes (from ventral to dorsal: the ventrolaterals [= second laterals], laterals [= first laterals], dorsolaterals, and paravertebrals; stripe names in brackets from Markezich et al. [1997]); dark-hued fields between the stripes; and pale-colored spots. One or more secondary, partial to complete, vertebral (= mid-dorsal) pale-colored stripes between the paravertebrals were also present in all specimens. Table 1. Data reductions for SVL (mean ± 1 SE, range, and number ) for adult females and males in samples of Cnemidophorus lemnsicatus from Miami-Dade County, FL and Isla de Utila, Honduras (only means and SE followed by all different superscripted letters are significantly different). Florida, USA Utila, Honduras Mean SVL Range n Mean SVL Range n All adult females 61.2 ± 1.49BC 54–64 (6) 60.6 ± 1.56C 56–68 (9) All adult males 71.0 ± 3.20A 61–88 (7) 67.9 ± 1.35AB 61–74 (10) 2009 B.P. Butterfield, J.B. Hauge, A. Flanagan, and J.M. Walker 47 We present tabular comparisons of meristic characters and ratios based on 16 specimens of C. lemniscatus from Miami-Dade County, FL, and 30 from Isla de Utila, Islas de la Bahía, Honduras, one of the nearest natural populations of the species, located over 1200 km southwest of Miami in the Caribbean Sea (Table 2, Appendix 1). Data were also available for over 500 specimens of C. lemniscatus from Guatemala, Honduras (including islands), and Panama in Central America, and Brazil, Colombia, Guyana, Suriname, and Venezuela in South America (J.M. Walker, unpubl. data). Qualitative characters of scutellation studied include position of each nostril with respect to a nasal suture, shape of the frontonasal scale, structure of the enlarged paracloacal scales (= anal spurs), and size of the posterior angle of the enlarged central preanal shield (see Burt 1931, Markezich et al. 1997). Meristic characters and ratios compared include the numbers of granules (scales) around mid-body (GAB), granules from occipital scales to base of tail (OR), ratio (x 100) of the granules around mid-body to granules from occipital scales to base of tail (GAB/OR), granules between the paravertebral stripes at mid-body (PV), ratio (x 100) of the granules between the paravertebral stripes at midbody to the granules around mid-body (PV/GAB), femoral pores combined (FP), subdigital lamellae of the longest toe of the left pes (SDL), circumorbital scales combined (COS), lateral supraocular granules combined (LSG), parietal scales (PS), mesoptychial scales (posteriormost transverse row of enlarged scales of the mesoptychium [MS]), scale rows anterior to the edge of the posterior gular fold (SAG; Burt 1931), scales bordering the outer edge of the parietal scales between the circumorbital series on each side (SBP), and interlabial scales between the chin shields and infralabials (ILS). Table 2. Data reductions (mean ± 1 SE and range) for 12 meristic characters, one of color pattern (PV), and two ratios useful in comparisons of samples (n) of Cnemidophorus lemnsicatus from Miami-Dade County, FL, and Isla de Utila, Honduras (see text for meaning of abbreviations; asterisk denotes that means for a character are significantly different). Florida, USA (n = 16) Utila, Honduras (n = 30) Character Mean Range Mean Range GAB* 106.2 ± 1.19 100–115 99.1 ± 0.71 93–109 OR* 205.1 ± 2.50 180–218 219.4 ± 1.83 201–239 FP* 39.9 ± 1.01 33–47 42.3 ± 0.41 38–48 SDL 31.1 ± 0.38 28–34 31.7 ± 0.12 31–33 COS* 10.5 ± 0.40 7–13 7.0 ± 0.19 4–9 LSG* 13.0 ± 0.88 7–23 10.1 ± 0.29 8–14 SO 8.1 ± 0.13 8–19 7.9 ± 0.07 7–8 PS* 5.4 ± 0.18 5–7 5.0 ± 0.14 5–6 MS* 11.9 ± 0.35 8–14 10.4 ± 0.22 8–14 SAG 2.3 ± 0.20 1–3 2.1 ± 0.12 1–3 SBP* 13.7 ± 0.28 12–16 12.1 ± 0.18 10–14 ILS* 6.6 ± 0.49 2–9 2.0 ± 0.00 2 PV* 14.9 ± 0.49 12–19 19.1 ± 0.30 14–22 PV/GAB* 14.1 ± 0.49 10.6–17.2 19.2 ± 0.29 15.1–22.4 GAB/OR* 51.7 ± 0.93 45.6–60.8 45.3 ± 0.41 41.4–51.5 48 Southeastern Naturalist Vol. 8, No. 1 We used JMP, v. 7 software (SAS Institute Inc., Cary, NC 1987–2007) to run ANOVA to generate a mean ± 1 SE and range of variation for SVL pooled by sex from each site, and for each univariate meristic character in the geographical samples of C. lemniscatus. Means were compared for statistical significance (P = 0.05) using Students t tests in JMP. Results Body size Sexual dimorphism in SVL was indicated in samples of C. lemniscatus from Miami-Dade County and Isla de Utila; males were significantly larger in mean SVL than females (Table 1). No significant differences in mean SVL were apparent by sex between the two samples (Table 1). The fact that the SVL of the largest Florida male examined (FLMH 131539; 88 mm) exceeded that of the largest preserved male examined from Isla de Utila (FLMNH 28379; 74 mm) was a sampling error is indicated by a male of 88 mm caught and released on the island by C.E. Montgomery (Truman State University, Kirksville, MO, pers. comm.). Punzo (2001) reported females (to 105 mm SVL ) and males (to 100 mm SVL) of C. lemniscatus from a site in Miami (= Hialeah); these are much larger maximum sizes for this species than we were able to confirm during this study (see Discussion). Reproduction Two female C. lemniscatus of 61 and 63 mm SVL (AUM 34112–34113) collected by B.P. Butterfield and J.B. Hauge at Virginia Gardens on 13 July 1996 contained two oviductal and two yolked ovarian eggs, respectively. Two females of 63 and 64 mm SVL (B.P. Butterfield, 1960–1961) collected at the same site on 20 May 1999 had two oviductal and three yolked ovarian eggs, respectively. Reproductive activity in Florida C. lemniscatus apparently began well before May in 1999. Among eight specimens collected by B.P. Butterfield and J.B. Hauge in Hialeah on 20 May 1999 were three recent hatchlings of 30, 30, and 29 mm SVL (B.P. Butterfield, 1967–1969), and two females of 54 and 62 mm SVL (B.P. Butterfield, 1962 and 1964) with one and three yolked ovarian eggs, respectively. We infer that embryonic development of these hatchlings began in March 1999. Based on the smallest gravid female of C. lemniscatus examined (54 mm SVL; collected 20 May 1999), Rainbow Whiptail in Florida mature in the first 12 months of life. This individual likely represented a late July–early August hatchling of the previous year. Conceivably, juveniles collected on 20 May 1999 could have attained reproductive size by September; however, egg production in such individuals of C. lemniscatus could be delayed until the following year. For the six gravid females of this species from Florida, there was no relationship between clutch size and SVL (r2 = 0.15). Color pattern Three Florida juveniles of C. lemniscatus, two females and one male (29.9 ± 1.0 mm SVL), had nine pale-colored longitudinal stripes and intervening 2009 B.P. Butterfield, J.B. Hauge, A. Flanagan, and J.M. Walker 49 dark brown fields. The white primary stripes on each side of the body include (ventral to dorsal) the ventrolateral, lateral, dorsolateral, and paravertebral (the two lower ones also extend anteriorly along the sides of the head). The vertebral or mid-dorsal, the darkest colored and least sharply-defined of the stripes (with only the slightest indication of an anterior bifurcation at the parietal scales), was positioned between the paravertebrals. The two lower primary stripes on each side (ventrolateral and lateral) differed from the two upper stripes (dorsolateral and paravertebral) in being irregular in outline, fused with spots, and ontogenetically variable. Distinct spots were present on the hind legs of each juvenile; some of these spots persisted through adulthood in both sexes. In each of the seven adult males of C. lemniscatus examined, the ventrolateral and lateral pairs of stripes existed as only vaguely discernable remnants, having the appearance of becoming faded and replaced by spots (Meshaka et al. 2004). Such adult males could be described as five-striped, although with age, the dorsolaterals become modified into series of spots in the manner of the ventrolaterals and laterals. The six adult females of C. lemniscatus from Florida, which in life were not as colorful as the seven males, showed less evidence of extensive ontogenetic change in color pattern (loss of the two lower pairs of stripes had not progressed as far as in the males). The fully metamorphosed color pattern of adult male C. lemniscatus in Florida (Meshaka et al. 2004; and on Isla de Utila (Gutsche 2005) underscores the basis of the common name, Rainbow Whiptail, and is indicative of the presence of this species, albeit limited, in the pet trade. Scutellation Diagnostic of the C. lemniscatus complex is a pointed scale lateral to the cloaca in males of all gonochoristic species (Burt 1931). Markezich et al. (1997) reported that males of C. lemniscatus have differently shaped paracloacal scales (= anal spurs) compared with males of C. gramivagus McCrystal and Dixon (Llanos Whiptail) and C. arenivagus Markezich, Cole, and Dessauer (Peninsula Whiptail). Florida males clearly possess the character state described for C. lemniscatus by Markezich et al. (1997): “... very broad-based spur tapering abruptly to a point near the apex...” In addition, specimens of Florida C. lemniscatus have each nostril opening more anterior than posterior to the nasal suture, a variably shaped frontonasal scale (that only occasionally could be considered hexagonal), and a central preanal shield with an obtuse posterior angle (i.e., only slightly greater than 90 degrees) (see Markezich et al. 1997). Meristic Characters Means for 15 univariate meristic characters between eight specimens of each sex in C. lemniscatus from Florida and between 14 males and 16 females of the species from Isla de Utila indicated an absence of sexual dimorphism. Comparison of pooled samples of males and females from each area revealed that only three characters, (SDL, SO, SAG) were not significantly different among localities (Table 2). However, means for 12 characters (GAB, OR, PV, PV/GAB, GAB/OR, FP, COS, LSG, PS, MS, SBP, ILS) were significantly different (Table 2). 50 Southeastern Naturalist Vol. 8, No. 1 Ecology This account is based on observations made by A. Flanagan in Hialeah, the most recent ones being 10 May and 2 August 2008. Although C. lemniscatus has been established and breeding in Hialeah at East 8th Avenue and East 16th Street for only about three years, the species has been present in surrounding areas (e.g., railroad right-of-way) for many years. In the vicinity of this address, this species is abundant in habitat consisting of patches of open-structured vegetation around buildings, residential dwellings, and swale, areas with varying amounts of litter/debris. Lizards utilize substrates consisting of soil, paved streets, and sidewalks. Evidence that C. lemniscatus is breeding at the site is based on observation of a range of size classes. Lizards persist there by adapting to use of man-made structures and associated microhabitats dominated by introduced plants in the absence of natural cover. Examples of types of structures frequented by lizards include hedge borders, fence rows with a variety plants, and any nook or cranny that provides retreats and/or shelter. Rainbow Whiptail can be observed basking on landscape timbers, sidewalks, concrete curbs, driveways, and/or decorative piles of rocks such as those occupied by lizards observed and/or photographed by A. Flanagan in May and August 2008. These observations in Hialeah indicated that these lizards are strictly terrestrial (they have not been seen in trees or on walls). Frequently used retreats by C. lemniscatus are either burrows of their own construction or spaces under concrete slabs for air conditioner compressors, sidewalks, and driveways. C. lemniscatus is well-established in Hialeah for several blocks in all directions from the East 8th Avenue site. Discussion Although Rainbow Whiptail is established in the literature as the common name of C. lemniscatus in Florida (Conant and Collins 1998, Meshaka et al. 2004), it cannot be assumed that all uses of it pertain to this species. For example, two respondents to our request for information on Florida C. lemniscatus via reported the presence of what they referred to as Rainbow Whiptail at two sites in Broward County. Further investigation of these reports conclusively revealed that one site is inhabited by native Aspidoscelis sexlineata (L.) (Six-lined Racerunner) and that the other is inhabited by the exotic species Ameiva ameiva (L.) (Giant Ameiva), both members of the Family Teiidae. Consequently, reports of Rainbow Whiptail in Florida should be accepted as C. lemniscatus only if they are vouchered by specimens or photographs of lizards from the sites in question. Lack of such vouchers made it impossible for us to resolve ambiguities raised by the SVL data for putative C. lemniscatus from Florida reported by Punzo (2001). He mentioned 100–123 mm SVL for the adult lizards used in a study of diet composition, but subsequently stated that adult males of 60–100 mm and adult females of 70–105 were used. Not only do we consider the upper size limits given by Punzo (2001) to be suspiciously large for C. lemniscatus, the lack of sexual dimorphism in Punzo’s sample (males 2009 B.P. Butterfield, J.B. Hauge, A. Flanagan, and J.M. Walker 51 expected to be larger) was also surprising. Unfortunately, all specimens of C. lemniscatus from that study were destroyed in a fire at the University of Tampa (F. Punzo, University of Tampa, FL, pers. comm.) prior to this study. Although the study site described by Punzo (2001) is situated only about 6 km from the Virginia Gardens site, 0.5 km from the Hialeah sites visited by B.P. Butterfield, and is the same as the site of origin for the individual of C. lemniscatus (KU 209859) posted on, we could not reconcile Punzo’s SVL measurements to known size data for this species from throughout its range (see Cole and Dessauer 1993, Colli et al. 2003, Markezich et al. 1997, this study). Of other taxa in the Family Teiidae established in Florida (Meshaka et al. 2004), only one species superficially resembles C. lemniscatus. Both native A. sexlineata (sensu Reeder et al. 2002) and Florida C. lemniscatus have a pale-striped dorsal pattern either during all (the former) or at certain (the latter) stages of ontogeny. However, A. sexlineata has an unspotted dorsal pattern and stripes throughout life (i.e., six primary stripes and vertebral = mid-dorsal; distinct ventrolaterals are lacking except on the neck). Juveniles of C. lemniscatus have eight white primary stripes (including distinct ventrolaterals), a mid-dorsal, and distinct spots; adults undergo extensive ontogenetic changes in dorsal pattern, with most stripes being replaced by spots in males and to a lesser extent in females. Syntopy between these species in Florida has not been reported. Using color pattern and meristic data, our analyses eliminated five of six species in the C. lemniscatus complex as the source of founders of this Florida adventive. The excluded species, countries of occurrence, and distance from Miami-Dade County to nearest population (based on Google Earth) are: parthenogenetic C. cryptus Cole and Dessauer (Icabaru Whiptail) (Brazil and Venezuela; 2960 km to Icarubu, Venezuela) and C. pseudolemniscatus Cole and Dessauer (Suriname Triploid Whiptail) (Suriname; 3600 km to Paramaribo) described by Cole and Dessauer (1993); gonochoristic C. gramivagus (Brazil, Colombia, and Venezuela; 2450 km to Apuré, Venezuela) described by McCrystal and Dixon (1987) and C. arenivagus (Venezuela; 2279 km to Paraguana Peninsula) described by Markezich et al. (1997); and gonochoristic C. nigricolor Peters (Black Racerunner) (several Caribbean islands of Venezuela; 2030 km to Los Roques Archipelago) diagnosed as a species by Walker and Rhoads (2003). There is no ambiguity pertaining to our assertion that neither of the parthenogenetic species in the C. lemniscatus complex is represented among the Florida specimens referenced herein, one of us also having examined more than 300 triploid C. pseudolemniscatus from Suriname and the type series of diploid C. cryptus from Venezuela for comparisons (J.M.Walker, unpubl. data). A breeding population of C. lemniscatus has existed in Florida for between >20 (Punzo 2001) and >40 (King and Krakauer 1966) years. All of the sites known for C. lemniscatus in Miami-Dade County are within the general vicinity of the Florida East Coast Railway. Habitats utilized by this species in Florida, like the one previously described (see Results), have enough open-structured vegetation and sufficient exposed pebbly/sandy soil 52 Southeastern Naturalist Vol. 8, No. 1 and artificial substrates for essential lizard activities (e.g., basking, foraging, mating, fl eeing, etc.), and structures that provide shelter for burrows and oviposition (e.g., debris, buildings, shrub rows, railroad ties, etc.). Based on information provided separately by K. Enge (Florida Fish and Wildlife Conservation Commission, Quincy, FL, pers. comm.) and R. St. Pierre (, pers. comm.), this species has achieved a very high density along the railway in Hialeah and in nearby neighborhoods. According to R. St. Pierre, C. lemniscatus likely has a more lengthy distribution along the railway than the 8.5-km stretch where it was actually observed by him. Moreover, he noted that the species has a continuous distribution between all of the sites listed herein for Miami-Dade County, where it occurs in areas from which native herpetofauna has been largely extirpated, an exception being Pantherophis (= Elaphe) guttatus (L.) (Corn Snake), which is a predator on this lizard. This anecdotal evidence on abundance, and the ca. 30 lizards observed by A. Flanagan in about 45 minutes on 2 August 2006, indicates that two attributes of C. lemniscatus in Florida, its high fecundity and early maturation, are the same ones advantageous to colonization ability also evident in most other established exotic amphibian and reptile species of Florida (Meshaka 2006, Meshaka et al. 2004). There is no way to be certain about the number of neotropical populations of C. lemniscatus from which whiptail lizards were imported to Florida; however, the uniformity in the fully metamorphosed color pattern of Floridian males of this species points to a single source. It was possible to eliminate some regional populations as the source of founders using the methodologies employed in this study. We infer that Florida specimens refl ect the diagnostic color pattern attributes (i.e., of juveniles and adults) and scalation characters (i.e., position of nostril and shapes of frontonasal scale, anal spurs, and central preanal shield) of founders. For the univariate meristic characters presented in Table 2, interspecific and intraspecific comparisons pertaining to samples of the C. lemniscatus complex yielded results for means and ranges of variation that are statistically significant or not, rather than present or absent (Colli et al. 2003, Markezich et al. 1997, Walker and Rhoads 2003). Thus, with the exceptions of the number of transverse rows of ventral scales (not included in Table 2), ILS, and SO, most univariate characters would be unreliable for use as predictors as the source of founders. Using the ILS character, we opine that Florida founders (mean 6.6 ± 0.49, range 2–9, n = 16) could not have been imported from Honduran islands Utila, Roatán, and Cayo Cochino Pequeño because those populations are characterized by one interlabial scale on each side. We also exclude populations of C. lemniscatus in Panama as the likely source of founders because they are characterized by unusual asymmetrical configurations of right and left supraocular scales (SO) and presence of 10 transverse rows of ventral scales compared to symmetrical 4-4 SO and eight rows of ventrals for Florida lizards. Using stripe characteristics, we concluded that founders of the Florida population of C. lemniscatus did not originate in Brazil, Guyana, or Suriname; all populations therein are characterized by either 10–11 stripes or a split vertebral stripe. Certainly, C. lemniscatus splendidus of the Peninsula de Paraguana, Venezuela, is excluded based 2009 B.P. Butterfield, J.B. Hauge, A. Flanagan, and J.M. Walker 53 on its almost completely blue dorsal coloration in adults. Thus, the most likely sources of founders for the Florida population of C. lemniscatus are populations in Venezuela, except as noted, Colombia, and some island in the Caribbean Sea. In turn, biochemical genetics would provide the best method to test our hypothesis concerning founders. Acknowledgments We are grateful to Ron St. Pierre for directing us to the Hialeah and Virginia Garden sites for C. lemniscatus. We thank Paul Bedsole, Joe Burgess, Kay Butterfield, and the Freed-Hardeman University Field Research Class for assistance in the collection of specimens. Walter E. Meshaka, Jr. provided especially helpful comments on the manuscript. We acknowledge the assistance of Joseph Collins for e-mailing a solicitation for information on C. lemniscatus in Florida to recipients of CNAH postings. This resulted in a helpful chain of responses pertaining to this species from Kevin Enge, Walter Meshaka, Jr., Carol Morgenstern, Ann Paterson, Trevor Persons, Ron and Stella St. Pierre, and Howard York. We are especially grateful to the following curators, collection managers, and individuals for either loans or transfers of specimens and photographs: Christy McCain and Mariko Kageyama, University of Colorado Museum, loans of specimens of parthenogenetic species in the C. lemniscatus complex from Suriname; Kenneth Krysko, University of Florida Museum of Natural History, loan of specimens from Islas de Utila and Roatán, Honduras; Charles Cole, American Museum of Natural History, loans of specimens from Guyana, Suriname, and Veneauela; Jose P. Rosado, Museum of Comparative Zoology, loan of specimens from Venezuela; and Chad Montgomery, then of University of Wisconsin-La Crosse, transfer of specimens and/or photographs of lizards from Panama and Honduras. Literature Cited Bartlett, R.D., and P. P. Bartlett. 1999. A field guide to Florida reptiles and amphibians. Gulf Publishing Company, Houston, TX. Burt, C.E. 1931. A study of the teiid lizards of the genus Cnemidophorus with special reference to their phylogenetic relationships. Bulletin of the United States National Museum 154:1–286. Cole, C.J., and H.C. Dessauer. 1993. Unisexual and bisexual whiptail lizards of the Cnemidophorus lemniscatus complex (Squamata: Teiidae) of the Guiana Region, South America, with descriptions of new species. American Museum Novitates 3081:1–30. Colli, G.R., G.C. Costa, A.A. Garda, K.A. Kopp, D.O. Mesquita, A.K. Péres, Jr., P.H. Valdujo, G.H.C. Vieira, and H.C. Wiederhecker. 2003. A critically endangered new species of Cnemidophorus (Squamata: Teiidae) from a Cerrado Enclave in southwestern Amazonia, Brazil. Herpetologica 59:76–89. Conant, R., and J. T. Collins. 1998. A field guide to reptiles and amphibians of eastern and central North America. Houghton and Miffl in Co., Boston, MA. Gutsche, A. 2005. Beobachtungen zu natürlichen Inkubationsbedingungen von Cnemidophorus lemniscatus (Linnaeus, 1758) (Sauria: Teiidae) auf der Isla de Utila, Honduras. Sauria 27:13–16. King, F.W., and T. Krakauer. 1966. The exotic herpetofauna of southeast Florida. Quarterly Journal of the Florida Academy of Sciences 29:144–154. 54 Southeastern Naturalist Vol. 8, No. 1 Markezich, A.L., C.J. Cole, and H.C. Dessauer. 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. American Museum Novitates 3207:1–60. McCrystal, H.K., and J.R. Dixon. 1987. A new species of Cnemidophorus (Sauria: Teiidae) from the Llanos of Colombia and Venezuela. Journal of Herpetology 21:245–254. Meshaka, W.E., Jr. 2006. An update on the list of Florida’s exotic amphibian and reptile species. Journal of Kansas Herpetology 19:16–17. Meshaka, W.E., Jr., B.P. Butterfield, and J.B. Hauge. 2004. The exotic amphibians and reptiles of Florida. Krieger Publishing Company, Malabar, FL. Montgomery, C.E., R.N. Reed, H.J. Shaw, S.M. Boback, and J.M. Walker. 2007. Distribution, habitat, size, and color pattern of Cnemidophorus lemniscatus (Sauria: Teiidae) on Cayo Cochino Pequeño, Honduras. Southwestern Naturalist 52:38–45. Peters, J.A., and R. Donoso-Barros. 1970. Catalogue of the Neotropical Squamata. Part II. Lizards and amphisbaenians. U. S. National Museum Bulletin 297:viii + 1–243. Punzo, F. 2001. Diet composition of the rainbow whiptail, Cnemidophorus lemniscatus (Sauria: Teiidae), from southern Florida. Herpetological Review 32:85–87. Reeder, T.W., C.J. Cole, and H.C. Dessauer. 2002. Phylogenetic relationships of whiptail lizards of the genus Cnemidophorus (Squamata: Teiidae): A test of monophyly, reevaluation of karyotypic evolution, and review of hybrid origins. American Museum Novitates 3365:1–61. Walker, J.M., and D.D. Rhoads. 2003. The black racerunner (Cnemidophorus nigricolor Peters, 1873) in Los Roques Archipelago, Venezuela. Herpetological Review 34:105–109. Wilson, L. D., and L. Porras. 1983. The Ecological Impact of Man on the South Florida Herpetofauna. University of Kansas Museum Natural History Special Publication No. 9. University Kansas Press, Lawrence, KS. 89 pp. Appendix 1. Sites of known occurrence for Cnemidophorus lemniscatus in Florida and specimens examined. USA: Florida: Miami-Dade County: [26 March 2002 (FLMNH 131480–131482, 131539)]; Virginia Gardens, Florida East Coast Railway between Northwest 39th Street and Northwest 38th Terrace [13 July 1996 (AUM 34112–34113, n = 2); 20 May 1999 (BPB 1960–1961, n = 2)]; Hialeah, Florida East Coast Railway in the vicinity of East 21st Street and East 11th Avenue.[20 May 1999 (BPB 1962–1969, n = 8)]. Hialeah, 1.3 km Southeast of East Coast Railway Building at Northwest 37th Avenue and 75th Street [7 March–30 August 1999 (n = 30: Punzo 2001); 17 July 1988 (Image of KU 209859, n = 1)]. Hialeah, East 8th Avenue and East 16th Street, 25o50'13.89"N, 80o15'56.17"W [10 May 2008 and 2 August 2008 (Images of a juvenile, a female, and several males by A. Flanagan)]. Honduras: Islas de la Bahía: Isla de Utila [13 August 1969 (FLMNH 28366– 28388, n = 23); 14 August 1969 (FLMNH 28430–28435, n = 6); 15 August 1969 (FLMNH 28436, 28444, n = 2); 17 August 1969 (FLMNH 28485, n = 1).