Observations on Nesting and Clutch Size in
Furcifer oustaleti (Oustalet’s Chameleon) in South Florida
Dustin Smith, Joy Vinci, Christopher V. Anderson, Jennifer Ketterlin Eckles, Frank Ridgley, and Frank J. Mazzotti
Southeastern Naturalist, Volume 15, Special Issue 8 (2016): 75–88
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Observations on Nesting and Clutch Size in
Furcifer oustaleti (Oustalet’s Chameleon) in South Florida
Dustin Smith1, 2,*, Joy Vinci3, Christopher V. Anderson4, 5,
Jennifer Ketterlin Eckles6, 7, Frank Ridgley1, and Frank J. Mazzotti3
Abstract - We studied an established population of Furcifer oustaleti (Oustalet’s Chameleon)
in southeastern Florida to understand aspects of reproductive biology in this
nonnative species. Reproduction of Oustalet’s Chameleon had not been documented in the
field in Florida, and limited information is available about its reproduction in its native
range. We conducted surveys from July 2011 to October 2012 in the Persea americana
(Avocado) grove where this species was introduced in Miami-Dade County, FL. During
these surveys, we removed more than 430 chameleons. We encountered 23 gravid females
from June to October. Mean clutch size was 42.3 eggs, and we recorded a new maximum
clutch size of 72 eggs. Utilizing radio-telemetry, we were able to track a gravid female to a
nest cavity, and herein describe the first Oustalet’s Chameleon nest in Florida. Our findings
suggest that management for eradication of the species should include ongoing surveys,
with removal efforts intensified from June to October, when females are known to be gravid.
Introduction
Florida has 56 established species of nonnative herpetofauna, more than any
other state in the US (Engeman et al. 2011, Krysko et al. 2011). Little is known
about the life history of many of these tropical and subtropical species within their
native ranges. However, much less is known about how they interact with the environment
in their introduced range. Significant resources are rarely spent learning
about introduced species until they become invasive and cause economic, ecological,
or human health or safety concerns; by then, eradication is usually expensive
and nearly impossible (Byers et al. 2002, Mehta et al. 2007, Pimentel et al. 2005).
Better understanding of the ecology of these species may improve the ability to
develop management actions to address potential impacts.
The number of introduced species of reptiles and amphibians in the state of Florida
continues to increase. Most introductions occur through anthropogenic means
such as cargo transport or the pet trade (Krysko et al. 2011,Wilson and Porras 1983).
Of the 56 species of established nonnative herpetofauna documented in Florida,
46 are lizards (Krysko et al. 2011), 2 of which are species of Chamaeleonidae:
1Conservation and Research Department, Zoo Miami, Miami, FL 33177. 2Current address
- North Carolina Zoological Park, Asheboro, NC 27205. 3Fort Lauderdale Research and Education
Center (FLREC), University of Florida, Davie, FL 33314. 4Department of Integrative
Biology, University of South Florida, Tampa, FL 33620. 5Current address - Department of
Biology, University of South Dakota, Vermillion, SD 57069. 6Florida Fish and Wildlife Conservation
Commission, Davie, FL 33314. 7Current address - FLREC, University of Florida,
Davie, FL 33314. *Corresponding author - Dustin.smith@nczoo.org.
Manuscript Editor: Will Selman
Everglades Invasive Species
2016 Southeastern Naturalist 15(Special Issue 8):75–88
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Chamaeleo calyptratus Duméril & Bibron (Veiled Chameleon; see Krysko et
al. 2004) and Furcifer oustaleti (Mocquard) (Oustalet’s Chameleon; see Gillette
et al. 2010). Other chameleon species have been introduced in Florida (e.g., Engeman
et al. 2005, Krysko et al. 2011, Rochford et al. 2013), but these are regarded
as either extirpated or representative of isolated escapees/releases that have not
formed established populations.
Oustalet’s Chameleon is the second largest extant species of chameleons, reaching
up to 68.5 cm total length (Glaw and Vences 1996, 2007; Nečas 2004). Although
the diet of most chameleons consists primarily of arthropods (Hofer et al. 2003,
Kraus et al. 2011), Oustalet’s Chameleon is known to eat small vertebrates including
birds (García and Vences 2002). Endemic to Madagascar, Oustalet’s Chameleon
naturally inhabits dry deciduous forests, wet montane savannahs, and primary forest
(Glaw and Vences 2007). It also thrives in agricultural lands and anthropogenically
altered landscapes (D’Cruze and Kumar 2011, Glaw and Vences 2007). This species
was previously introduced into Kenya, where its current status is unknown (Nečas
2004; Spawls et al. 2002, 2014; Tilbury 2010). At least 16 y ago, Oustalet’s Chameleon
was introduced to a single 48-ha Persea americana Mill. (Avocado) grove
in southwestern Miami-Dade County, FL. All size classes including neonates have
been observed at this site (Gillette et al. 2010; D. Smith et al., unpubl. data).
Although the introduction of Oustalet’s Chameleon in southeastern Florida
has been described (Gillette et al. 2010, Krysko et al. 2011), there has been no
documentation of reproduction in this population. A better understanding of the
ecology and life history of the Oustalet’s Chameleon may be critical for developing
effective management strategies for this nonnative species. Here, we report on
chronology of gravid females, nest-site characteristics, clutch size, egg characteristics,
and incubation period for individuals collected in southeastern Florida.
Materials and Methods
Surveys
Our study site was a privately owned 48-ha Avocado grove located in Florida
City, FL (25.433ºN 80.501ºW). Three sections of the grove were divided into 400 m
x 400 m plots. Although the grove was predominantly composed of Avocado trees,
we also found chameleons on Schinus terebinthifolius Raddi (Brazilian Pepper),
Neyraudia reynaudiana (Kunth) Keng ex A.S. Hitchc. (Burma Reed), Parthenocissus
quinquefolia (L.) Planch. (Virginia Creeper), Parthenium hysterophorus L.
(Santa Maria Feverfew), and other nonnative and native plant species. The grove is
bordered by a row of Roystonea regia (Kunth) O.F. Cook (Royal Palm) and a road
to the south, an Avocado grove to the west, and roads on the northern and eastern
boundaries. The Navy Wells Pineland Preserve, a natural area consisting primarily
of pine-rockland habitat managed by the Miami-Dade County Parks, Recreation,
and Open Spaces Department, is adjacent to the northern perimeter road (Fig. 1).
We conducted nighttime visual surveys (n = 27) with flashlights from 12 July
2011 to 15 October 2012 to remove Oustalet’s Chameleons and determine their
reproductive characteristics; additional data was collected for related studies. We
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performed surveys throughout the grove and in a section of Navy Wells Pineland
Preserve to the north. Surveys began after sunset and lasted 3–4 h, depending on
the number of individuals surveying and whether the survey was constrained or
unconstrained. We conducted 9 unconstrained surveys throughout the study site
from July to October 2011. Following these surveys, we conducted 19 distanceconstrained
surveys along standard routes from 20 October 2011 to 15 October
2012. Each distance-constrained survey consisted of walking 1 of 4 selected areas
(1 in each of 3 plots in the Avocado grove, and 1 plot included a portion of the
Navy Wells preserve) of ~12,000 m2, 1–2 times each month. Although we did not
conduct formal surveys in adjacent areas, informal visual-encounter surveys were
conducted in July 2011 in all adjacent, potential sites.
When Oustalet’s Chameleons were encountered, we recorded the location (GPS
coordinates), size, sex, and reproductive status of females. We captured and removed
all specimens unless they were too high in the trees to safely capture. We
determined gravidity of females based on body size and coloration at the time of
collection (see Cuadrado 2000). Gravid females have a distended abdomen and are
green with colors that are not observed throughout the rest of the year, including
Figure. 1. Map showing three 400 m x 400 m Avocado grove plots and the adjacent Navy
Wells Pineland Preserve. We established 1 survey route in each of these 4 areas.
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white, red, or orange patterns (Fig. 2a). Non-gravid females have a more-slender
abdomen and are green with a less colorful background pattern (Fig. 2b).
Figure. 2. Visual differences between a (A) gravid and (B) non-gravid female Oustalet’s
Chameleon in Persea americana (Avocado) trees. Note the distended abdomen and
different coloration in the gravid female compared to the non-gravid female. The transmitter
attachment saddle using leader line is also shown in panel A, including location
and connections (arrow).
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Clutch data and incubation
Gravid females were kept individually in approximately 60 cm x 60 cm x 90
cm-tall nylon-mesh-sided enclosures with live plants, full spectrum and basking
lighting on a 12-h light cycle, and daily misting. We provided nesting buckets with
25–40 cm of a sand and soil mixture as oviposition sites. Following oviposition,
we weighed and measured (length x width) a subsample of the eggs (n = 149) from
15 clutches. We used this subset of clutch data because females were distributed
among different facilities and incomplete data were collected at some locations.
Clutches from 3 females, as well as a clutch from 1 other female collected from the
same site prior to the start of our surveys, were artificially incubated in deli-cup
containers filled midway with a 1:1 water:vermiculite mixture by weight. No additional
moisture was added to the media during the incubation period. We placed the
containers inside a dark closet and allowed the eggs to incubate at ambient laboratory
temperature (19–23 ºC average daily temperature range) to test the resilience
of eggs to a broad range of incubation temperatures. We used linear regression
analysis (α = 0.05) and the 2-tailed Pearson product-moment correlation coefficient
to test for correlations between female mass and clutch mass and between snout–
vent length (SVL) and clutch size. Statistical analyses were performed in R, and
means are reported as ± standard deviation (SD).
Radio-telemetry
We captured a single gravid female on 15 October 2012, affixed a VHF radio
transmitter (BD-2, Holohil, ON, Canada) on its dorsal ridge, and radio-tracked her
to determine oviposition location (Fig. 2a). The transmitter was attached to a saddle,
created using a 2.5-mm-vinyl plastic-coated leader line. We secured the transmitter
saddle over the pelvic region, with the line wrapped both cranial and caudal to
her hind limbs; leader-line ends were attached to one another using a stainless steel
barrel-crimp with its edges wrapped in tape (Fig. 2a). We released the specimen at the
collection site on 16 October 2012 at ~1610 h. Thereafter, we tracked it daily using a
handheld receiver (R-1000, Communications Specialists, Inc., Orange, CA) and Yagi
antenna. We used a handheld weather meter (Kestrel Meter 3000, Nielsen-Kellerman,
Boothwyn, PA) to measure the nest temperature and humidity.
Results
Chronology of reproduction
We collected 431 Oustalet’s Chameleons (78 males, 81 females, and 272 unsexable
juveniles) during surveys from July 2011 to October 2012. We captured a total
of 23 gravid females during June (n = 6; 100% of females captured), July (n = 6;
75%), August (n = 6; 75%), September (n = 3; 21%), and October (n = 2; 50%).
We captured 11 gravid females from July to October 2011, and 12 from June to
October 2012. An additional gravid female used in this study was collected opportunistically
in May 2011 prior to the start of structured surveys. During July 2011 to
October 2012, all collected gravid females (n = 24) oviposited during June (n = 3),
July (n = 4), August (n = 10), September (n = 2), and October (n = 5). Only a single
clutch was recorded for each female included in this study.
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Clutch data
We collected egg and clutch data from 24 gravid female Oustalet’s Chameleons. A
total of 1014 eggs (mean = 42 ± 11.76; range = 17–72) were laid. Gravid females averaged
164 ± 1.53 mm SVL (n = 14) with a mean post-capture mass of 124.0 ± 35.8g
(n = 12) (Table 1). Mean egg dimensions among 15 clutches were 16.7 ± 1.3 mm x
10 ± 0.88 mm (n = 149). Artificially incubated clutches (n = 4) began hatching after
285–354 days, and all viable eggs (59–93% of each clutch) hatched within 21 days of
the first hatching from their respective clutches. Clutch size relative to female SVL
(n = 14) was not significant (R² = 0.20; P = 0.06; Fig. 3), while clutch mass and female
mass (n = 9) were significantly correlated (R² = 0.49; P = 0.04; Fig. 4).
In situ nesting
We released the female chameleon outfitted with a radio transmitter at the point
of capture and began tracking on 17 October 2013. She stayed in the same tree for
6 d, and then we tracked her to the base of an Avocado tree 42 m west of the initial
capture and release site. We found 2 open cavities in the substrate at the base of
the Avocado tree where she was tracked. Avocado trees at this site are planted in
large, elevated soil mounds because of the solid limestone substrate in southeastern
Florida. The opening of the larger of the 2 cavities was on the southern side
of the Avocado tree facing southward. The cavity entrance was 9 cm wide and the
egg closest to the nest-chamber entrance was at a depth of ~42 cm from the cavity
opening. Upon further excavation, we found the chamber to be ~6 cm wide and 23
cm long. The chamber contained 58 eggs scattered throughout; some were partially
buried by substrate. The clutch weighed 60.2 g, with a mean egg length of 17.2 ±
0.93 mm (n = 13, range = 16–19 mm) and width of 10.4 ± 0.51 mm (n = 13, range
= 10–11 mm). We measured the temperature and relative humidity of the interior
chamber to be 27.2 °C and 94.9%, respectively. Ambient air temperature was 27.3
Table 1. Individual, clutch, and egg morphometric data for gravid female Oustalet’s Chameleons collected
in Florida 2011–2012. n = sample size for egg dimension data.
Capture Post- Oviposition Clutch Mean egg
month SVL capture month Clutch weight length width
ID and year (mm) mass (g) and year size (g) (mm) (mm) n
67 July 2011 162 105 July 2011 32 - - - -
69 July 2011 158 75 August 2011 39 36 16 10 10
95 July 2011 164 110 August 2011 49 30 15 8 10
118 August 2011 160 120 August 2011 47 - - - -
122 August 2011 170 110 August 2011 35 36 18 9 10
127 August 2011 165 110 August 2011 34 26 16 9 10
149 October 2011 140 100 October 2011 39 38 16 11 10
310 June 2012 145 92 June 2012 42 30 17 11 6
312 June 2012 195 188 June 2012 62 76 16 11 10
327 June 2012 165 130 July 2012 49 49 15 9 10
328 June 2012 182 170 July 2012 44 50 19 10 10
397 August 2012 145 - August 2012 41 46 17 10 10
410 August 2012 165 - August 2012 35 39 18 10 10
427 October 2012 184 178 October 2012 58 - - - -
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°C with 94.9% relative humidity. We could not determine if the female dug these
cavities or utilized some that were already present. After data collection, we removed
and discarded the eggs from this clutch to preempt hatching.
Discussion
Our study provides the first documentation of natural reproduction in Oustalet’s
Chameleon in Florida, as well as a description of a nesting site and a new maximum
clutch size. Little information has been published on chameleon natural history (see
Herrmann and Herrmann 2005, Measey et al. 2014, Reaney et al. 2012), especially
related to reproduction and oviposition in wild populations. Published data suggest
that oviposition for chameleons occurs during the summer months, when rains are
at or near their peak (Andrews and Karsten 2010, Cuadrado 1999, Herrmann and
Herrmann 2005, Karsten et al. 2008, Reaney et al. 2012; Table 2). Our data support
this premise, including our observed oviposition by wild-collected gravid female
Oustalet’s Chameleons that oviposited from June to October, which are the summer
and rainy-season months in Florida. However, stress from capture and handling or
husbandry variations could have caused early or delayed oviposition.
Figure 3. Linear regressions of clutch size vs. female SVL (R² = 0.20; P = 0.06). Dots represent
individual clutch data from separate females. The dashed line depicts a non-significant
relationship.
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Figure 4. Linear regressions of clutch mass vs. female mass (R² = 0.49; P = 0.04). Dots
represent individual clutch data from separate females. The solid line depicts a significant
correlation.
Chameleons are known to reproduce seasonally or continuously throughout
the year, depending on biotic and abiotic factors, with seasonally reproducing
chameleons often ovipositing prior to winter and having long incubation lengths
that include diapause (Measey et al. 2014). Chameleons may have adapted and
extended the timing of breeding and nesting seasons in regions with stable temperatures
and highly seasonal rainfall (Shine and Brown 2008). Avoidance of nest
and offspring predation may also contribute to an extended nesting period (Jackson
1988). The reproductive phenology for Oustalet’s Chameleon is unknown, but our
data, including the pre-winter oviposition and an extended incubation length, suggest
that there may be an extended, seasonal reproduction season in southeastern
Florida. Although the incubation lengths we observed may be biased because of
captive-temperature stability or lower incubation-temperatures, incubation time
is extended for the species. Environmental factors contributing to this extended,
seasonal reproduction season may be the subtropical climate with moderate yearround
temperatures and seasonal precipitation. As in other lizard species, a major
contributing factor to seasonal reproduction in Oustalet’s Chameleon may be availability
of prey for offspring (Fitch 1982).
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Clutch sizes vary greatly among chameleon species. Clutch sizes tend to increase
with increasing SVL (Andrews and Karsten 2010), but this relationship
can vary both interspecifically and intraspecifically (Table 2; e.g., Andrews et
al. 2008, Burrage 1973, Nečas 2004, Reaney et al. 2012). Unlike patterns among
some other chameleon species, such as Chamaeleo chamaeleon (L.) (Common
Chameleon) and interspecific relationships (Andrews and Karsten 2010, Cuadrado
1998), we did not observe a correlation between clutch size and female SVL
(Fig. 3). This difference may be due to the more limited range of body sizes in our
analysis relative to other studies, small sample size, or the influence of an outlier
(see Fig. 3). In addition, we found a positive correlation between female mass and
clutch mass (Fig. 4). The results of both our clutch size vs. female SVL and clutch
mass vs. female mass correlations may be strengthened by additional sampling,
particularly among larger females.
Oustalet’s Chameleon was previously known to have a maximum clutch size of 61
eggs (Brygoo 1971, Glaw and Vences 2007, Nečas 2004, Spawls et al. 2002), however
2 females in our study laid clutches of 62 and 72 eggs, respectively. Incubation
time for this species at a constant 28 ºC is known to range from 210 to 280 d (Nečas
2004), but we found that clutches can incubate at a much lower temperature range
(19–23 ºC) and still remain viable. However, incubation at lower temperatures leads
to considerably longer incubation times to hatching (285–354 d), ~75 d longer than
reported by Nečas (2004). These findings illustrate a high reproductive potential for
this species, and egg resilience to a wide range of incubation temperatures.
Utilizing radio telemetry to detect nesting locations of nonnative species has
been successful in southeastern Florida. In addition to our study, other radio-telemetry
studies have been able to find nesting locations for other invasive reptiles in
Florida including Salvator merianae A.M.C. Duméril & Bibron (Argentine Black
and White Tegu; Pernas et al. 2012) and Python molurus bivittatus Kuhl (Burmese
Python; Snow et al. 2007). Based on our tracking efforts with Oustalet’s Chameleon
and the temporal presence of gravid females, oviposition likely occurs from June
to October. Additional studies are needed to confirm in situ locations of nests to
determine if Oustalet’s Chameleon uses locations for oviposition sites other than
Avocado tree mounds.
This population of Oustalet’s Chameleon in southeastern Florida was intentionally
released in this privately owned Avocado grove (Gillette et al. 2010), possibly
for the same reason the Trioceros jacksonii xantholophus (Eason et al.) (Jackson’s
Chameleon) release locations were chosen in Hawaii. Jackson’s Chameleons were
released near Mangifera indica L. (Mango) and Psidium guajava L. (Guava) trees
because the fruit attracts fruit pests that are also prey for neonates (Kraus et al.
2011). Jackson’s Chameleons were then spread throughout the island and were
eventually sold in the pet trade. It is unclear if Oustalet’s Chameleon and other
chameleon species are being spread in a similar manner in southeastern Florida,
although it seems likely (Edwards et al. 2014). Agricultural lands in this area provide
fruit that attracts prey, which would be immediately encountered by neonates
hatching at the base of fruit trees. Most natural areas of Miami-Dade County have
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Table 2. Interspecific comparison of reproductive traits and asso ciated ecological factors of 6 oviparous chameleon species.
Female Oviposition Hatching
SVL Clutch size months Incubation months
Species Location (mm) Range (mean) (season) length (season) References
Chamaeleo chamaeleon L. Southern 87–135 6–28 (12) Sep–Oct 10.5–12 mo Aug–Nov Andrews et al. 2008,
(Common Chameleon) Europe (summer/ (summer/rainy) Cuadrado 1999
early rainy)
Trioceros montium (Buccholz) Southwestern 65–90 3–12 (6.5) Feb–Jun ~3.5 mo (in lab) ~May–July Hermann and Hermann
(Cameroon Sailfin Chameleon) Cameroon (summer/ (summer/rainy) 2005
early rainy)
Chamaeleo dilepis Leach Southern >80 mm 19–74 (44.2) Nov–May - - Reaney et al. 2012
(Common Flap-necked Chameleon) Africa (summer/
early rainy)
Furcifer labordi (Grandidier) Madagascar - - Jan–Mar 8–9 mo Nov–Dec Karsten et al. 2008
(Laborde’s Chameleon) (mid-late summer/ (early summer/
mid-late rainy) early rainy)
Chamaeleo namaquensis Smith Namibia 90–143 6–22 (11.8) May–Oct 91–112 d Sep–Jan Burrage 1973
(Namaqua Chameleon) (winter–spring/ (spring–summer)
rainy)
Furcifer oustaleti Mocquard Southeastern 145–195 17–72 (24) Jun–Oct 285–354 d Jul–Oct This study
(Oustalet’s Chameleon) Florida (summer/rainy) (summer–
early fall)
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a solid limestone substrate (Hoffmeister et al. 1967) in which it would be difficult
for Oustalet’s Chameleon to excavate a nest. The disturbed and enhanced soil
mounds and irrigation systems in Avocado groves provide adequate nesting sites
for chameleons and provide proper soil, moisture, and humidity throughout the
year. This combination likely provides Oustalet’s Chameleon a greater chance
for survival, reproductive success, and population persistence compared to native
habitats. Because of these factors, disturbed areas, such as agricultural or urban areas
may be suitable for some non-native lizards. The most widespread, introduced
chameleon in Florida is the Veiled Chameleon. It is established in at least 3 counties
(Enge 2008, Gillette and Krysko 2012, Krysko et al. 2004, Meshaka 2011) where
it thrives in agricultural areas and anthropogenically altered habitats similar to the
one reported herein. Although we are not aware of chameleons that have colonized
any natural areas in Florida, there have been other non-native reptiles that thrive
in natural areas. In southeastern Florida, the Argentine Black and White Tegu was
first observed in an agricultural area, and has been seen in a variety of disturbed and
natural areas (Pernas et al. 2012). Some of these sites also have disturbed mounds
of soil, which may provide suitable nesting sites and contribute to persistence of
new populations.
It is unclear if Oustalet’s Chameleon has a negative impact on native species, but
this species shows similar characteristics to other established nonnative species. Lizards
are the most successful nonnative reptile taxa in Florida, and they share many
traits including high fecundity and early maturation (Meshaka 2011). Successful
nonnative species also tend to have widespread native ranges, abundant populations,
and adapt well to climates outside of their natural range (Sakai et al. 2001,
Williamson and Fitter 1996). Oustalet’s Chameleon is fecund, having clutch sizes
of 17–72 eggs with offspring becoming sexually mature in 1 y (Glaw and Vences
2007; D. Smith et al., unpubl. data), and occurs in a variety of habitats throughout
much of Madagascar (Glaw and Vences 2007). In 1 survey of a dry deciduous forest
in northern Madagascar, Oustalet’s Chameleon was the most common reptile encountered
(Labanowski and Lowin 2011). Based on climate data from 2005 to 2015
at the Antsiranana Airport weather station in northern Madagascar, the mean annual
temperature there is 25.4 ºC (range = 12.2–32.4 ºC). This temperature is very similar
for the same range of dates in southeastern Florida at the Miami International Airport
weather station where the mean annual temperature was 25.1 ºC (range = 1.6–35.5
ºC); the range is wider and the minimum temperature is lower. We assume that this
species has adapted to the climate of southeastern Florida, as it has been at this Avocado
grove for more than 10 y (Gillette et al. 2010).
Knowledge gained from our study can aid in the management of Oustalet’s Chameleon
and other introduced species. Removal of reproductive individuals is a key
component in the eradication of an invasive species (Bomford and O’Brien 1995).
Based on knowledge of the reproductive season, removal efforts can be intensified
during that time. Because we encountered gravid females from June to October
and the majority of females oviposited in August, removal during these months,
with efforts peaking by the end of July, would be the most effective at reducing
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population growth. Although removal of the reproductive population is crucial, it is
as important to diligently study invasive species during the removal process to learn
about their biology. It is also imperative to create a management plan for removal,
and consistently survey until the population is eradicated.
Acknowledgments
This project was supported by Miami-Dade Parks, Recreation, and Open Spaces; University
of Florida; Florida Fish and Wildlife Conservation Commission; and the Everglades
Cooperative Invasive Species Management Area. We thank all of the volunteers who participated
in the fieldwork, especially Venetia Briggs-Gonzalez, Robin Bijlani, Rafael Crespo,
Michelle Curtis, Seth Farris, Sergio Gonzalez, Kevin Kopf, Mike Rochford, Sara Williams,
and Ryan Zach. We are grateful to Jessi Krebs, Cayle Pearson, and Ryan O’Shea for their
contribution of data, Dr. Krystal Tolley for valuable information regarding transmitter
placement on chameleons, and Nell Allen for assistance with the manuscript. We also thank
Grove Services, Inc., for allowing us access to the grove for this study. Lastly, we appreciate
feedback on the manuscript provided by Kevin Enge, Kelly Irick, Kristen Sommers, and 2
anonymous reviewers. Approval for this study was obtained from the Zoo Miami Animal
Care and Use Committee.
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