Journal of Thermal Biology 38 (2013) 41–46

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Journal of Thermal Biology

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Influences of sex, ontogeny and body size on the thermal ecology

of Liolaemus lutzae (Squamata, Liolaemidae) in a restinga remnant
in southeastern Brazil
Thiago Maia-Carneiro n, Carlos Frederico Duarte Rocha
Departamento de Ecologia, Universidade do Estado do Rio de Janeiro, Rua Sa~ o Francisco Xavier 524, CEP 20550-013, Rio de Janeiro, Brazil

a r t i c l e i n f o abstract

Article history: Variations in body temperature (Tb) of lizards can be partially explained by intrinsic factors such as sex,
Received 3 April 2012 ontogeny and body size. Liolaemus lutzae is a lizard species restricted to restingas in the Brazilian coast
Accepted 19 October 2012 in the state of Rio de Janeiro. Herein, we studied sexual dimorphism and influences of sex, ontogeny,
Available online 1 November 2012
and body size to the Tb of L. lutzae. Adult males were larger than adult females, probably due to both
Keywords: intersexual selection and intra-sexual selection. There was intersexual difference in lizards’ Tb (males
Atlantic rainforest hotter than females), but Tb did not differ after factored out for the effects of body size. The mean Tb of
Body mass juvenile lizards was higher than that of adults after factored out for the effect of body mass. It is
Body temperature possible that adults may have excluded juveniles from microhabitats with better thermal regimes. Also,
Sexual dimorphism
this might have occurred due to requirements of juveniles to maintain high growth rates. Forage
Snout–vent length
searching for prey by juveniles also exposes them to high environmental temperatures. Juveniles also
Thermal inertia
may have higher Tb than co-specific adults (relative to body mass) to favor prey capture. In absolute
values, adult lizards tended to use microhabitats with lower temperatures than that used by juveniles,
possibly to avoid risks of overheating and death. Body temperature and snout–vent length were
positively related, as well as body temperature and body mass, presumably caused by the thermal
inertia of the bodies (trend of a body to resist to changes in its temperature). Intrinsic factors such as
sex, ontogeny and body size can affect the thermal ecology of L. lutzae, despite coastal habitat features
to which they are exposed also influences the body temperature of active lizards in restinga habitats.
& 2012 Elsevier Ltd. All rights reserved.

1. Introduction in beach habitat of restingas, might offer risks of overheating for

Some lizard species can have variations in body temperature
partially explained by intrinsic factors such as sex and ontogeny
(Magnusson, 1993; Carothers et al., 1998) and body size (snout–vent
length and body mass, Powell and Russell, 1985; Stevenson, 1985;
Carothers et al., 1997). Behavioral adjustments can also have
important roles for body temperature regulation by active lizards
(Castilla et al., 1999; Rocha et al., 2009a). Furthermore, extrinsic
factors such as wind intensity (Maia-Carneiro, unpublished data)
and variation of environmental temperatures (Rocha, 1995a;
Vrcibradic and Rocha, 1995; Verrastro and Bujes, 1998; Rocha
et al., 2009b) also influence the thermoregulation by these animals,
and might cause changes in the body temperature of active
individuals. Indeed, elevated environmental temperatures, such as
n
Corresponding author. Tel.: þ55 2188923679.
E-mail addresses: thiagomaianc@gmail.com (T. Maia-Carneiro),
cfdrocha@uerj.br (C.F.D. Rocha).
lizards which may result in death (Mosauer, 1936).
Liolaemus lutzae Mertens, 1938 (Liolaemidae) is a lizard
species restricted to habitats of restinga in the Brazilian coast in
the state of Rio de Janeiro, with geographic distribution of
approximately 200 km between the Restinga da Marambaia
(231040 S, 431520 W), in the municipality of Rio de Janeiro, and
the restinga da Praia do Peró (221510 S, 411590 W), in the munici-
pality of Cabo Frio (Vanzolini and Ab’Saber, 1968; Rocha et al.,
2009c,d). This species occur in the habitat of halophilous–
psammophilous-reptant vegetation near to the sea in restingas
of the state (Rocha et al., 2009d; Maia-Carneiro and Rocha, in
press), a habitat under intense pressure due to anthropogenic
activities (Rocha et al., 2007, 2009d). This is true especially
concerning the area and the surroundings of where this study
was conducted (restinga of Praia Grande), which is under higher
anthropogenic pressure comparing to other restingas in the
region (Rocha et al., 2009e).
Information regarding thermal ecology and factors that can
influence this biological aspect of lizards is relevant to understand
the relationships between the species and the environment, and

0306-4565/$ - see front matter & 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.jtherbio.2012.10.004
42 T. Maia-Carneiro, C.F.D. Rocha / Journal of Thermal Biology 38 (2013) 41–46
to elucidate some components of its Natural History. This is
especially important whether is one of the lizard species in the
world recently indicated as having a great probability of extinc-
tion in the next few decades due to its thermal niche alteration
caused by climatic changes, such as L. lutzae (Sinervo et al., 2010).
Herein, we investigated the importance of intrinsic factors (sex,
ontogeny, and body size) and of thermal environments for
regulation of body temperature by active L. lutzae in a beach
habitat.
2. Materials and methods
2.1. Study area
Field sampling was conducted in a stretch of restinga in Praia
Grande (221 570 S, 421 020 W), municipality of Arraial do Cabo, state
of Rio de Janeiro, southeastern Brazil, near to the Área de Protec- a~ o
Ambiental da Massambaba. Restingas are habitats characterized
by soils with high salt concentration and a predominance of
herbaceous and shrubby vegetation in a sandy substrate (Suguio
and Tessler, 1984). The climate in the Área de Protec- a~ o Ambiental
da Massambaba is seasonal, with mean annual precipitation of
approximately 800 mm (most precipitation concentrated in the
summer) and mean annual temperature of 25 1C, varying between
12 1C and 36 1C (Scarano, 2002).
2.2. Data collection
Data were collected in June, July, and September (dry season),
and in November and December (rainy season) of 2011 through
visual encounter surveys (Crump and Scott, 1994) constrained by
time. We delimited a study area measuring 60 m of width by
500 m of length, totalizing approximately 3.0 ha of restinga
examined. Visual encounter surveys (30 min) were performed
inside the delimited area at each hour interval between 0600 h
and 1800 h, totalizing 4680 min of sample effort (2880 min in the
dry season and 1800 min in the rainy season). When an individual
of L. lutzae was sighted, were made capture attempts by hand.
At the time of collection we measured the body temperature (Tb)
(precision of 0.2 1C) of each captured lizard utilizing a Schultheis
quick-reading cloacal thermometer. We only considered the body
temperatures obtained within a maximal interval of 30 s after the
first sight of the lizard. We identified the sex of each lizard (Rocha,
1996a), recorded their snout–vent length (SVL) (measured with
caliper with precision of 0.1 mm) and their body mass (measured
with a dynamometer Pesolas with precision of 0.25 g). Additionally,
we measured the substrate temperature (Ts) in the local where the
lizard was first sighted and the air temperature (Ta) 1 cm above the
substrate in the same point using the Schultheis thermometer
(precision of 0.2 1C). We considered these microhabitat tempera-
tures as measurements of the thermal environment which the
lizards experienced.
2.3. Data analyses
To evaluate differences in SVL, body mass, and Tb between
male and female adult lizards we performed two-way multi-
variate analysis of variance (MANOVA). To analyze occurrence of
intersexual differences in body mass removing the effect of SVL
we made analysis of covariance (ANCOVA) (Zar, 1999). To eval-
uate whether the SVL, body mass, and Tb of juveniles and adults
differed we used MANOVA. To test for differences in Tb between
adult males and adult females and between juveniles and adults
of L. lutzae at each season factoring out the effects of SVL and of
body mass, we performed ANCOVA (Zar, 1999). To test for
differences in microhabitat temperatures (air and substrate)
experienced by juveniles and adults we made one-way analyses
of variance (ANOVA). We made simple regression analyses and
multiple regression analyses (Zar, 1999) between the body
temperatures and the body size to evaluate the importance of
SVL and of body mass for regulation of Tb by L. lutzae.
The ontogenetic stage (juvenile or adult) was accessed by
consulting the minimum size at maturity known for males and
females of L. lutzae (Rocha, 1992). We performed statistical
analyses separated per season considering the occurrence of
seasonal climatic differences (Scarano, 2002), which could affect
the Tb of L. lutzae (Rocha, 1995a; Maia-Carneiro, unpublished data).
Actually, microhabitat temperatures recorded at the time of collec-
tion of data of this study had higher values during the rainy season
compared to the dry season (Maia-Carneiro, unpublished data).
3. Results
With regard to adults L. lutzae, the SVL, the body mass, and the
Tb differed considering sex (MANOVA, Wilks’ Lambda ¼0.401,
F3,102 ¼50.789, Po0.0001), season (MANOVA, Wilks’ Lambda¼
0.044, F3,102 ¼744.756, Po0.0001), and the interaction between
these predictor factors [sex  season (MANOVA, Wilks’
Lambda¼0.733, F3,102 ¼12.368, Po0.0001)]. Adult males were
longer, heavier, and had higher Tb than adult females
(Tables 1 and 2), but the sexes did not significantly differ in body
mass after factored out for the effect of the SVL (ANCOVA,
R2 ¼0.089, F2,105 ¼0.041, P¼0.840). At the dry season, there was
intersexual difference in Tb (ANOVA, R2 ¼ 0.111, F1,36 ¼4.476,
P¼0.041), with males being comparatively hotter than females
(Table 2). At the rainy season, we did not detected intersexual
differences in Tb (ANOVA, F1,68 ¼3.469, P¼0.067) (Table 2).
At both seasons, there was no differences in Tb between sexes
(dry season: ANCOVA, F3,34 ¼0.137, P¼0.714; rainy season:
ANCOVA, F3,66 ¼1.604, P¼0.210) after factoring out the effect of
SVL (dry season: F3,34 ¼2.225, P¼0.145; rainy season: F3,66 ¼
2.474, P ¼0.121) and of body mass (dry season: F3,34 ¼1.954,
¼0.171; rainy season: F3,66 ¼2.427, P¼ 0.124).
The SVL, the body mass, and the Tb differed either considering
age class (MANOVA, Wilks’ Lambda ¼0.612, F3,215 ¼ 45.368,
Po0.0001), season (MANOVA, Wilks’ Lambda¼ 0.254, F3,215 ¼
210.811, P o0.0001), or the interaction between these predictors
[age class  season (MANOVA, Wilks’ Lambda¼0.810, F3,215 ¼
16.776, Po0.0001)]. Juveniles tended to be smaller, lighter, and
colder than adults (Table 2). At the dry season, the Tb of juvenile
lizards was proportionally higher than that of adults (ANCOVA,
R2 ¼0.280, F3,129 ¼5.214, P¼0.024), with body mass explaining an
additional portion of the variation in the Tb (F3,129 ¼6.661,
P¼0.011), which did not occur with the SVL (F3,129 ¼0.000,
P¼0.999). At the rainy season, there was no difference in Tb
Table 1
Number of individuals (N), arithmetic mean and one standard deviation (SD), and
range of snout–vent length (in mm) and of body mass (in g) of males and females
L. lutzae sampled at both dry and rainy seasons (pooled data) in a remnant of
restinga in Praia Grande, municipality of Arraial do Cabo, state of Rio de Janeiro, in
Brazilian east coast.
Body variable N Mean7 SD Range
Snout–vent length (mm)
Males 43 67.6 75.3 61.0–78.9
Females 65 56.5 73.8 51.5–75.5
Body mass (g)
Males 43 7.9 72.1 4.5–13.7
Females 65 4.7 71.1 3.2–11.0
 T. Maia-Carneiro, C.F.D. Rocha / Journal of Thermal Biology 38 (2013) 41–46 43

Table 2
Number of individuals (N), arithmetic mean and one standard deviation (SD), and range of snout–vent length (SVL, in mm), body mass (in g) and body
temperature (in 1C) of adult males, adult females, adultsa, juveniles, and pooled data of all individuals of L. lutzae sampled at the dry and rainy seasons in a
remnant of restinga in Praia Grande, municipality of Arraial do Cabo, state of Rio de Janeiro, in Brazilian east coast.

Body variable Dry season Rainy season

N Mean7 SD Range N Mean 7SD Range

SVL (mm)
Adult males 15 65.87 5.3 61.1–75.4 28 68.5 7 5.2 61.0–78.9
Adult females 23 55.27 3.3 51.5–62.7 42 57.2 7 3.9 51.6–75.5
Adultsa 38 59.37 6.6 51.5–75.4 70 61.7 7 7.1 51.6–78.9
Juveniles 95 43.97 6.9 32.0–58.9 18 51.4 7 4.5 45.8–60.3
All individuals 133 48.37 9.8 32.0–75.4 88 59.7 7 7.9 45.8–78.9

Body mass (g)

Adult males 15 7.87 2.6 4.5–13.7 28 8.07 1.8 5.2–13.5
Adult females 23 4.67 1.0 3.2–7.2 42 4.77 1.2 3.5–11.0
Adultsa 38 5.97 2.4 3.3–13.8 70 6.07 2.2 3.5–13.5
Juveniles 95 2.57 1.4 0.5–10.3 18 3.47 1.0 2.0–6.0
All individuals 133 3.57 2.3 0.5–13.7 88 5.57 2.3 2.0–13.5

Body temperature (1C)

Adult males 15 33.27 1.8 28.4–35.4 28 33.6 7 2.6 25.4–37.2
Adult females 23 31.47 2.8 24.0–37.2 42 32.2 7 3.3 23.2–37.0
Adultsa 38 32.17 2.6 24.0–37.2 70 32.7 7 3.1 23.2–37.2
Juveniles 95 30.87 3.5 23.4–38.0 18 31.1 7 4.2 23.2–36.4
All individuals 133 31.27 3.3 23.4–38.0 88 32.4 7 3.4 23.2–37.0

a
Pooled both males and females.

Table 3
Number of individuals (N), arithmetic mean and one standard deviation (SD), and range of substrate temperature (in 1C) and of air
temperature (in 1C) of juveniles and adults L. lutzae sampled at the dry and rainy seasons in a remnant of restinga in Praia Grande,
municipality of Arraial do Cabo, state of Rio de Janeiro, in Brazilian east coast.

Microhabitat temperatures Dry season Rainy season

N Mean7 SD Range N Mean7 SD Range

Substrate temperature
Juveniles 95 31.2 7 4.9 1C 20.2–43.0 18 35.3 7 6.2 22.6–47.6
Adults 38 31.1 7 4.6 22.8–46.6 70 34.1 7 5.6 22.0–48.0

Air temperature
Juveniles 95 28.5 7 3.6 22.4–38.0 18 30.2 7 2.9 23.6–37.2
Adults 38 27.8 7 2.6 24.0–36.2 70 29.8 7 2.6 22.6–35.4

between adult and juvenile lizards active in the habitat (ANCOVA,
F3,84 ¼ 1.026, P¼0.314) after removed for the effect of both SVL
(F3,84 ¼0.307, P ¼0.581) and body mass (F3,84 ¼0.664, P¼0.418).
At both seasons there were no differences between juveniles and
adults in Ts of microhabitats which they used (dry season:
ANOVA, F1,131 ¼0.023, P¼0.879; rainy season: ANOVA,
F1,86 ¼ 0.607, P ¼0.438) (Table 3). The Ta of microhabitats used
by juveniles and adults did not differ at both dry season (ANOVA,
F1,131 ¼ 0.976, P¼ 0.325) and rainy season (ANOVA, F1,86 ¼0.446,
P¼0.506, Table 3).
At the dry season, there were positive relationships between Tb
and SVL (simple regression analysis, R2 ¼0.202, F1,131 ¼32.759,
Po0.0001) (Fig. 1) and between Tb and body mass (simple regres-
sion analysis, R2 ¼0.244, F1,131 ¼42.326, Po0.0001) (Fig. 2, Table 1).
At the rainy season, Tb and SVL (simple regression analysis,
F1,86 ¼2.147, P¼0.147) and Tb and body mass (simple regression
analysis, F1,131 ¼2.994, P¼ 0.087) were not related (Table 1). At the
dry season, only body mass (P¼0.004) explained an additional
portion of the Tb of active lizards (multiple regression analysis,
R2 ¼0.251, F2,130 ¼21.758, Po0.0001) after the effect of other
intrinsic variable (SVL; P¼0.287) was removed. At the rainy season,
both the SVL (P¼0.561) and the body mass (P¼0.284) did not
explained an additional portion of the Tb of active L. lutzae (multiple
regression analysis, F2,85 ¼1.656, P¼0.197).
4. Discussion
Evaluation of our data revealed that L. lutzae in the restinga of
Praia Grande are sexually dimorphic in body size (SVL and body
mass), with males reaching larger sizes than females, similar to
that found in a population in the restinga of Barra de Maricá (state
of Rio de Janeiro) (Rocha, 1996a). The larger body size of male
lizards may be favored in situations of choice by females (inter-
sexual selection) due to higher quality of their territories or in
agonistic interactions during disputes for territories between
males (intra-sexual selection acting among males) (Vitt and
Cooper, 1985; Rocha, 1992, 1996a; Lappin et al., 2006; Lailvaux
et al., 2004; Herrel et al., 2007), with larger males having more
access to females. Data from the home range of L. lutzae in
another restinga habitat (Barra de Maricá), revealed that the
home range of adult males slightly overlapped, whereas the home
range of one adult male had the home range from one to three
females associated to it (Rocha, 1999). This suggests exclusion
among males, whose tend to defend exclusive areas (territories)
to have access to females (Rocha, 1999). The sexual dimorphism
of L. lutzae probably reflects both the result of egg investment by
females (Rocha, 1992, 1995b) and intra-sexual selection related to
territorial behavior by males (Rocha, 1996a, 1999). As individuals
of L. lutzae behave as opportunist foragers (Rocha, 1998) appear
44 T. Maia-Carneiro, C.F.D. Rocha / Journal of Thermal Biology 38 (2013) 41–46
Fig. 1. Relationship between activity body temperature (in 1C, N¼ 133) of lizards
Liolaemus lutzae and the snout–vent length (in mm, N ¼133) during the dry season
in the municipality of Arraial do Cabo, state of Rio de Janeiro, southeastern Brazil.
Fig. 2. Relationship between activity body temperature (in 1C, N¼ 133) of lizards
Liolaemus lutzae and the body mass (in g, in logarithm, N¼133) during the dry season
in the municipality of Arraial do Cabo, state of Rio de Janeiro, southeastern Brazil.
that competition for food resources is not a selective force that
conducted to sexual dimorphism in body size of L. lutzae in Barra
de Maricá (Rocha, 1996a) nor in the studied population of Praia
Grande.
Analysis of our data revealed that the dissimilarities in
observed values of Tb found for the sexes was caused by the
relative larger body size of males compared to females, and its
properties of thermal inertia (trend of a body to resist to changes
in its temperature – see below). The nearest geographically co-
generic lizard Liolaemus occipitalis, from southern Brazil, also did
not have intersexual differences in their Tb (Bujes and Verrastro,
2006) in an ecologically similar environment to that we studied in
Rio de Janeiro (restinga sand beach habitat). However, males
Liolaemus nigroviridis Müller and Hellmich, 1932 (Liolaemidae) in
the Andes can vary in their Tb due to differential behavior and
body mass, which did not occur with females (Carothers et al.,
1998). The authors argued that this occurs due to the behavior of
acquisition and maintenance of territories that can expose adult
males L. nigroviridis to high environmental temperatures, which
increases their Tb (Carothers et al., 1998). A similar event might
have occurred with males L. lutzae in Praia Grande, for which the
exposition to high environmental temperatures because of differ-
ential territorial behavior (Rocha, 1999) may potentially increase
their body temperature. Evaluation of data from thermal biology
studies with 56 desert lizard species from three continents
(Africa, Australia and North America) revealed that males and
females of a given species have similar body temperatures and
thermal biologies (Huey and Pianka, 2007), though few species
can sexually differ in Tb (Lailvaux et al., 2003; Huey and Pianka,
2007).
In this study, L. lutzae differed ontogenetically in SVL, body
mass, and Tb, with juveniles being smaller, lighter, and having
mean Tb relatively higher than adults after factored out for the
effect of body mass. As the thermal environment of a micro-
habitat is an environmental resource, juveniles would be more
exposed to higher environmental temperatures due to exclusion
by co-specific lizards from microhabitats with more adequate
thermal regimes. The home range of juveniles L. lutzae was
significantly smaller than that of adult males but not differing
from that of adult females (Rocha, 1999). This author suggested
that those features of home range size could be explained, at least
in part, by differences in body size among juveniles and male
adults, which in turn would promote some differences in moving
rate along habitat: (1) juveniles of L. lutzae initiate dispersion just
after hatching, in order to establish their own home ranges, which
probably results in an increase in their movement rate;
(2) juveniles of L. lutzae are mainly carnivores (adults are
omnivorous but mainly consuming plants) feeding essentially
on terrestrial arthropods (Rocha, 1998) in a habitat with relatively
low arthropod availability (Rocha, 1996b), forcing them to move
over sand beach to obtain appropriated amount of their arthropod
prey. These differences in movement along habitat together with
differences in microhabitats frequented could explain, at least in
part, the observed differences in Tb between juveniles and adults
L. lutzae. However, the temperatures of microhabitats used did
not statistically differ with regard to ontogeny; although in terms
of absolute values, juvenile lizards tended to use thermal envir-
onments with higher temperatures than did adults in both
seasons.
We should also consider the role of ontogeny (instead competi-
tive exclusion) as a factor affecting the Tb of L. lutzae. In this way,
juveniles might have higher thermal requirements to maintain high
growth rates (Sinervo and Adolph, 1989; Rocha, 1995b; Rhen and
Lang, 1999), reaching proportionally higher body temperatures
relative to body sizes. Moreover, juveniles L. lutzae are predomi-
nantly carnivorous, opposed to adults that are mainly herbivorous
(Rocha 1998). Forage searching for prey in a habitat with high
environmental temperatures such as restingas exposes the lizards to
high temperatures, potentially increasing their Tb. As at the rainy
season can have greater prey availability than at the dry season for L.
lutzae (Rocha, 1996b), presumably they might spend less time
foraging and therefore be less exposed to high environmental
temperatures. Juveniles also may have higher Tb than co-specific
adults (relative to body sizes) to favor prey capture during the dry
season, when prey availability can be lower (Rocha, 1996b). Reduced
levels of solar radiation can affect the efficiency of predatory
behaviors decreasing sprint velocity while foraging, decreasing the
distance of forage and, hence, of the likelihood of encounter with
 T. Maia-Carneiro, C.F.D. Rocha / Journal of Thermal Biology 38 (2013) 41–46
potential prey, and decreasing the efficiency of search and manip-
ulation of prey (Avery et al., 1982). In this case, juveniles L. lutzae
might allocate more time on thermoregulation to avoid the effects of
reduced levels of solar radiation during the dry season compared to
the rainy season. Higher body temperatures also may increase flight
success by lizards from potential predators (Hertz et al., 1982; Rocha
and Bergallo 1990), besides affect food digestion by different reptile
species (Van Damme et al., 1991 – lizard; Wang et al., 2003 – snake;
Pafilis et al., 2007 – lizards).
Evaluation of our results of both seasons revealed that
temperatures of occupied microhabitats did not significantly
differ ontogenetically for L. lutzae in Praia Grande. However, in
terms of absolute values, adults tended to use microhabitats with
lower temperatures than did juveniles. This may have occurred
because the use by adults of thermal environments which
diminish risks associated with overheating. Large-sized lizards
have lower surface/volume relationships and as consequences:
(1) heat up and cool down slower; (2) have increased thermal
inertia; and (3) are less affected by convective cooling (Porter and
Gates, 1969; Carothers et al., 1997). Therefore, large lizards
maintain body temperatures easier than small lizards and
because of that may have a greater risk of overheating in hot
open environments (Porter and James, 1979; Paulissen, 1988a).
This might be true also in restinga beach habitats, where tem-
peratures can reach values which may impose death risk for
lizards (see Mosauer, 1936; Rocha, 1988; Table 3). Therefore,
adults L. lutzae may increase their Tb due to exposition to high
environmental temperatures and retain heat for longer time
periods, resulting in high body temperatures along day, and likely
increased risk of overheating and death, mainly in the rainy
season when environmental temperatures are higher. Moreover,
large lizards have greater thermal inertia and thus should be able
to stay in microhabitats exposed to low temperatures for longer
periods of time than smaller co-specifics because large lizards
would not experience a rapid drop in Tb.
The relationships between Tb and SVL and between Tb and
body mass (with larger lizards tending to have higher body
temperatures) presumably were caused by the thermal inertia
of the bodies. Thermal inertia is the trend of a body to resist to
changes in its temperature, where larger bodies have capacity of
heat retention for longer time periods than smaller bodies due to
its reduced surface/volume relationships (Carothers et al., 1997;
Garrick, 2008). The higher thermal inertia of larger lizards (with
longer SVL and heavier body mass) allows maintenance of the Tb
for a longer time, whereas the relatively lower thermal inertia of
smaller lizards causes easier heat loss from them. Such as at the
dry season in Praia Grande only body mass explained an addi-
tional portion of the variation in Tb of active lizards, the changes
in the Tb appeared to be more dependent of body mass than of
SVL. In general, current available information reveal a lack of
relation between Tb and body size for lizard species in different
families (e.g. Huey, 1982; Pianka, 1986; Paulissen, 1988b;
Magnusson, 1993; Rivera-Vélez and Lewis, 1994; Rocha, 1994;
Rocha and Vrcibradic, 1996; Carothers et al., 1998; Vrcibradic and
Rocha, 1998; Menezes et al., 2000; Kiefer et al., 2005; Menezes
and Rocha, 2011), which suggest that each species has mean
Tb in activity suitable to perform its different ecological and
physiological activities regardless of body size or age class.
4.1. Conclusions
L. lutzae is a sexually dimorphic lizard species in which males
achieve larger body sizes than females. These results are sugges-
tive of intersexual and intra-sexual selection acting on males and
females of this species. The variability in the Tb of L. luzae can
reflect inherent characteristics of individuals and thermal
45
patterns of local environments. Intrinsic factors such as sex,
ontogenetic stage and body size of individuals can be important
influences to their thermal biology, despite characteristics of the
coastal habitat to which they are exposed (open area subject to
high environmental temperatures and intense and constant wind
regimes – Rocha, 1995a; Maia-Carneiro, unpublished data) can
also influence the body temperature of active lizards of this
species in restinga habitats. The knowledge about processes
of thermal ecology of lizards is relevant for understanding of
relationships between individuals and the environments where
they occur. This might be of particular relevance if is a lizard
species under great probability of extinction due to its thermal
niche alteration such as L. lutzae (Sinervo et al., 2010). Such
knowledge and understanding can be used for definition of
strategies aiming conserving remnants of natural environments
(habitats where L. lutzae occurs) and therefore also natural
processes either physical, chemical, and in different levels of
biological organization.
Acknowledgments
This study was supported by research grants from the Con-
selho Nacional de Desenvolvimento Cientı́fico e Tecnológico–
CNPq to C.F.D. Rocha (processes 304791/2010-5 and 470265/
2010-8). C.F.D. Rocha also received financing from the Fundac- a~ o
de Amparo a Pesquisa do Estado do Rio de Janeiro–FAPERJ
through the ‘‘Programa Cientistas do Nosso Estado’’ (process
E-26/102.404.2009). T. Maia-Carneiro received an M.Sc. scholar-
ship from the Coordenac- a~ o de Aperfeic- oamento de Pessoal de
Nı́vel Superior (CAPES), and currently receives Ph.D. scholarship
from CAPES. We also thank to T.A. Dorigo who kindly revised the
manuscript and C.C. Siqueira who’s helped with statistical
analyses.
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