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
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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.
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 - email@example.com.
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 www.cnah.org (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.
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. ); 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
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
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.
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).
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).
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; www.cnah.org) 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.
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).
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
50 Southeastern Naturalist Vol. 8, No. 1
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.
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 www.cnah.org 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 www.cnah.org, 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
(firstname.lastname@example.org, 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.
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.
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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).