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Thermoregulation by a population of Aspidoscelis calidipes from Apatzingan, Michoacan, Mexico.

The body temperature of lizards is a reflection of their overall ecology including use of habitat, foraging tactics, nutritional state, and times of activity. Therefore, knowledge of thermal relationships can aid in constructing an integrative overview of the general ecology of species of lizards. Reptilian body temperature is affected by phylogeny (Casas-Andreu and Gurrola-Hidalgo, 1993) and environment (Pianka, 1970; Gillis, 1991; Bueter and Haas, 2008). Reptiles use behavioral mechanisms to regulate body temperature for optimal functioning of cellular and organismal processes (Bartholomew, 1982). Behavioral thermoregulation includes changes in posture (Kearney, 2001), activity periods (Grant and Dunham, 1988), and selection of microhabitat (Adolph, 1990). Because thermoregulation in reptiles is basically behavioral, they tend to explore the temporal and spatial variability of the thermal environment, which promotes a close relationship between body temperature of the organism and temperature of the environment (Mathies and Andrews, 1995). Among squamate reptiles, some of the highest active body temperatures ([T.sub.b]) occur in the Teiidae family, which suggests that this characteristic may have arisen early in this evolutionary history (Vitt and Pianka, 2004). Aspidoscelis calidipes (Duellman, 1960) belongs to the sexlineata group, which is the most diverse clade within the genus Aspidoscelis (Reeder et al., 2002). The focal species of our study is an endemic lizard of Mexico with an altitudinal range between 200 and 600 m above sea level. It inhabits the Tepalcatepec Valley characterized by xerophilous scrubland in the Balsas River Basin in the state of Michoacan. We analyze thermoregulation in A. calidipes and describe the relationship of body temperature with temperature of air and substrate.

This study was conducted in Las Cabas, Apatzingan, SW Michoacan, Mexico (18[degrees]33029.6"N, 102[degrees]59027.3"W, at an altitude of 343 m). Climate of the study area is tropical with rains in summer and dry steppe. Mean annual temperature and precipitation are 39.8[degrees]C and 924 mm, respectively. Vegetation includes tropical dry forest, tropical spiny forest, and mixed forest (Instituto Nacional de Estadistica y Geografia, http//:www.inegi.gob.mx). We collected 52 lizards during 1000-1800 h in April and June 2008 and January and May 2009. Active lizards were captured by noosing and were returned to their habitat after data were collected. We measured cloacal body ([T.sub.b]), substrate ([T.sub.s] on the site where the lizard was first observed), and air ([T.sub.a] 5 cm above the ground) temperatures with a quick-reading thermometer (model 4D2672, Miller and Weber, Inc., New York; 0.0-50.0 [+ or -] 0.2[degrees]C). Care was taken to prevent temperature from being influenced by handling, and all data from lizards requiring extensive effort to capture were excluded from records of temperature. To avoid errors, we only analyzed data collected within 10 s after capture (Lee, 1979; Gillis, 1991). In addition to [T.sub.b], we recorded snout-vent length (SVL) with an electronic caliper to the nearest 0.1 mm and measured body mass with a Pesola scale (Pesola AG, Baar, Switzerland) to the nearest 0.2 g. Sex was identified by examining the sexually dimorphic pattern of color and scales.

The assumptions of normality and homogeneity of variances were met for all analyses with Shapiro-Wilk and Bartlett tests, respectively. Statistical analyses were conducted with Statistica 9 Software (StatSoft, Tulsa, Oklahoma). We calculated residual from the relationship of body temperature to SVL to produce body-temperature adjusted variables that maintained variation of extrinsic factors and minimized confounding effects of size. If we did not find a significant relationship, we used original data to make the respective statistical analyses. We performed a Mann-Whitney U test to determine whether or not there are differences in body temperature, SVL, and body mass by sex. Additionally, we performed a Spearman rank correlation ([r.sub.s]) to examine the relationship among body, air, and substrate temperatures. Significance was accepted at P < 0.05. Means are presented plus and minus standard deviation.

Aspidoscelis calidipes had a mean SVL of 58.46 [+ or -] 9.14 mm (range of 41-76 mm) and a mean weight of 4.30 [+ or -] 2.44 g (range of 1.65-12.0 g). Overall mean [T.sub.b] was 40.44 [+ or -] 2.55[degrees]C (range of 32.6-44.6[degrees]C). There was no significant relationship between SVL and body temperature of A. calidipes (y = 36.669 + 0.064 SVL, [r.sup.2] = 0.053, F = 2.814, P = 0.099). Therefore, we used original data to compare body temperature by sex. We did not find differences between sexes in SVL (U = 0.165, P = 0.868) and body mass (U = 0.566, P = 0.571). We also did not find significant differences in [T.sub.b] between sexes (U = -1.478, P = 0.139). Mean [T.sub.b] of males was 40.26 [+ or -] 2.14[degrees]C (range of 37-44.6[degrees]C), while that for females was 41.36 [+ or -] 2.72[degrees]C (range of 34.4-44.0[degrees]C). Mean air temperature in Las Canas was 37.35 [+ or -] 3.81[degrees]C with a range of 31-45[degrees]C, and mean substrate temperature was 40.13 [+ or -] 4.55[degrees]C with a range of 31-50[degrees]C. Body temperatures of A. calidipes had a significant positive relationship with air ([r.sub.s] = 0.599, P < 0.0001) and substrate temperatures ([r.sub.s] = 0.542, P < 0.0001).

Our results showed that mean [T.sub.b] of A. calidipes was highest in comparison to other species of Aspidoscelis (Table 1). This high body temperature could be related with high environmental temperatures recorded during the study period. We did not find differences in body temperature between sexes. This demonstrates that males and females thermoregulate similarly, probably due to similar microhabitats (open forest), activity periods, and feeding habits as in other diurnal lizards (Guizado-Rodriguez et al., 2011).

Body temperature in A. calidipes was affected by [T.sub.a] and [T.sub.s]. Environmental temperatures frequently affect the [T.sub.b] of species of Aspidoscelis (Vitt et al., 1993; Guizado-Rodriguez and Casas-Andreu, 2007; Navarro-Garcia et al., 2008; Woolrich-Pifia et al., 2011). However, other species of Aspidoscelis do not show a relationship between body and environmental temperatures. Preferably, the extent to which a lizard thermoregulates would be determined by comparing body temperatures with operative environmental temperatures estimated using biophysical models (Heath, 1964; Hertz, 1992). Unfortunately, the appropriate data are not always available, as in this study. Therefore, additional fieldwork supplemented by studies of physiological tolerance-limits and thermal preferences in the laboratory would be relevant to complement the information presented here. In conclusion, the high body temperature and the little recorded variation between individuals of different sex of A. calidipes suggest that this characteristic in strongly influenced by phylogeny.

We are grateful to Instituto de Biologia and Posgrado en Ciencias Biologicas of Universidad Nacional Autonoma de Mexico. We also thank Consejo Nacional de Ciencia y Tecnologia for graduate-study scholarships to MAGR (CONACYT no. 165072). We thank C. Duifhuis-Rivera, U. O. GarciaVazquez, J. Maceda-Cruz, M. Fernandez-Aguilar, M. A. GarciaMorelos, C. A. Mendoza-Palmero, M. Ramirez-Sanchez, and G. Hernandez-Flores for their help in the field.

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Submitted 12 March 2013. Acceptance recommended by Associate Editor Felipe de Jesus Rodriguez Romero 25 May 2013.

MARTHA ANAHI GUIZADO-RODRIGUEZ, * LORENA REYES-VAQUERO, AND GUSTAVO CASAS-ANDREU

Departamento de Biologia, Universidad Autonoma Metropolitana Iztapalapa, AP 55-35, Avenida San Rafael Atlixco No. 186 Colonia Vicentina, Delegacion Iztapalapa, Moxico Distrito Federal, CP 09340 (MAGR)

Departamento de Biotecnologia, Centro de Desarrollo de Productos Bioticos, Instituto Politocnico Nacional, Carretera Yautepec-Jojutla, Km 6, Calle CEPROBI No. 8, Colonia San Isidro, Yautepec, Morelos, Moxico CP 62731 (LRV)

Departamento de Zoologia, Instituto de Biologia, Universidad Nacional Autonoma de Moxico, Circuito Exterior s/n, AP 70-153, Moxico Distrito Federal 04510 (GCA)

* Correspondent: anahigurz@gmail.com
TABLE 1--Mean body temperatures (Tb) for species of Aspidoscelis.

Species            Tb ([degrees]C)             Source

A. calidipes          32.6-44.6                   Present study
A. ceralbensis          40.7           Soule, 1963; Brattstrom,
                                                           1965
A. communis             36.2                   Casas-Andreu and
                                          Gurrola-Hidalgo, 1993
A. deppii             40.0-42.5          Kennedy, 1968; Vitt et
                                                      al., 1993
A. dixoni               39.89               Walker et al., 1991
A. exsanguis          38.5-39.9           Medica, 1967; Schall,
                                             1977; Bowker, 1993
A. flagellicauda        39.9                      Stevens, 1980
A. gularis            38.2-41.0       Brattstrom, 1965; Schall,
                                      1977; Bowker and Johnson,
                                        1980; Paulissen et al.,
                                          1989; Paulissen, 1999
A. guttata              40.4                      Kennedy, 1968
A. hyperythra           39.9           Soule, 1963; Brattstrom,
                                                           1965
A. inornata           38.6-40.2           Medica, 1967; Schall,
                                      1977; Bowker and Johnson,
                                                           1980
A. lineatissima      31.14-38.44               Casas-Andreu and
                                         Gurrola-Hidalgo, 1993;
                                          Guizado-Rodriguez and
                                            Casas-Andreu, 2007;
                                           Navarro et al., 2008
A. neomexicana          39.0                       Medica, 1967
A. parvisocia           37.16        Woolrich-Pida et al., 2011
A. sackii               38.29        Woolrich-Pida et al., 2011
A. sexlineata         36.6-41.0      Bogert, 1949; Fitch, 1958;
                                        Brattstrom, 1965; Witz,
                                                           2001
A. tesselata          40.1-42.0       Bogert, 1949; Brattstrom,
                                             1965; Schall, 1977
A. tigris             38.9-40.4               Brattstrom, 1965;
                                      Cunningham, 1966; Medica,
                                            1967; Pianka, 1970;
                                                  Schall, 1977;
                                     Lemos-Espinal et al., 1997
A. uniparens            38.6           Bowker and Johnson, 1980
A. velox                38.7                       Bowker, 1993
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Title Annotation:NOTES
Author:Guizado-Rodriguez, Martha Anahi; Reyes-Vaquero, Lorena; Casas-Andreu, Gustavo
Publication:Southwestern Naturalist
Article Type:Report
Geographic Code:1MEX
Date:Mar 1, 2014
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