ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah Acta Herpetologica 8(2): 123-128, 2013 Comparative ecophysiology of two sympatric lizards. Laying the groundwork for mechanistic distribution models Enrique García-Muñoz1,2,3*, Miguel Angel Carretero1 1 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de Vairão, 4485-661 Vairão, Portugal. *Corresponding author. E-mail: engamu@gmail.com 2 CESAM, Centro de Estudos de Ambiente o do Mar, Universidade de Aveiro, Campus Universitário de Santiago, 3810-193 Aveiro, Portugal 3 Departamento de Biología Animal, Biología Vegetal y Ecología. Campus de las Lagunillas, s/n. Universidad de Jaén. 23071 Jaén, Spain Submitted on 2013, 26th July; revised on 2013, 3rd December; accepted on 4th December. Abstract. Distribution modelling usually makes inferences correlating species presence and environmental variables but does not take biotic relations into account. Alternative approaches based on a mechanistic understanding of bio- logical processes are now being applied. Regarding lacertid lizards, physiological traits such as preferred body temper- ature (Tp) are well known to correlate with several physiological optima. Much less is known about their water ecol- ogy although body temperature and evaporative water loss (Wl) may trade-off. Two saxicolous lacertids, Algyroides marchi and Podarcis hispanica ss are sympatric in the Subbetic Mountains (SE Spain) were they can be found in synto- py. Previous distribution modelling indicates the first species is associated with mountains, low temperatures; high precipitation and forest cover whereas the second one is more generalistic. Here, we perform two ecophysiological tests with both species: a Tp experiment in thermal gradient and a Wl experiment in sealed chambers. Although both species attained similar body temperatures, A. marchi lost more water and more uniformly in time than P. hispanica ss that displayed an apparent response to dehydration. These results suggest that water loss rather temperature is crucial to explain the distribution patterns of A. marchi in relation to P. hispanica ss, the former risking dehydration in dry areas no matter what temperature is. Ecophysiological traits represent a promising tool to build future mechanistic models for (lacertid) lizards. Additionally, the implications for their biogeography and conservation are discussed. Keywords. Water loss, preferred body temperature, mechanistic models, Algyroides marchi, Podarcis. INTRODUCTION Thermal ecology is central in ecophysiological studies on lizards, and lacertids are not an exception to this rule (Castilla et al., 1999). Past decades gave rise to a plethora of detailed studies describing the proximate mechanisms linking temperature to physiology, life history, and behav- iour (Angilletta, 2010). Preferred body temperature in the absence of thermoregulatory constraints (Tp) constitutes an important trait in lizard studies correlating with sev- eral physiological optima (Huey and Bennet, 1987; Bauw- ens et al., 1995; Angilletta et al., 2002; Carretero, 2008a). In contrast, lizard water ecology has been neglected. Although protocols for testing water loss rate (Wl) already exist (Lillywhite, 2006) and even when morphological proxis have been found (Caslbeek et al., 2006) no compar- ative results are really available. Nevertheless, preliminary evidence suggests that body temperature and evapora- tive water loss may trade-off (Bruce, 1990; Bowker, 1993). Those species capable to reduce Wl while remaining mobile have the best opportunities for a precise control of temperature (Tracy, 1976; Buttemer, 1990). An animal with low Wl may move to regulate its temperature pre- cisely without experiencing the cooling and dehydrating effects of evaporation (Tracy and Christian, 2005). Nonetheless, both types of variables (Tp and Wl) are essential to develop mechanistic models of potential dis- tribution of species (Adolph and Porter, 1993; Fei et al., 124 Enrique García-Muñoz, Miguel Angel Carretero 2012). Species distribution modeling is currently domi- nated by correlation analyses. That is, to infer the factors restricting the range of an organism, its presence is cor- related to the environmental variables prevailing in its range. This procedure produces good results (Peterson and Kitchell, 2001; Martínez-Freiria et al., 2008; Santos et al., 2009), especially when the species have low disper- sal abilities (Sillero et al., 2009). However, other external variables can be involved namely, biotic factors which can keep species absent from environmentally suitable areas (Kearney and Porter, 2004). Among these factors, compet- itive interactions between ecologically similar species may restrict species ranges (Begon, 2006). Even if species inter- actions can be modeled based on correlation approaches (Costa et al., 2008), the determination of physiological traits may provide solid evidence (Porter et al., 2002,). Podarcis and Algyroides are two Mediterranean lac- ertid genera with highly contrasting richness and dis- tribution patterns. While 21 species of Podarcis wall liz- ards are currently recognized, most of them with broad continuous distributions (Arnold et al., 2007), Algyroides consist of only four species with separate relict ranges (Harris et al., 1999). Among them, Algyroides marchi is the most saxicolous species restricted to a very small range in the Subbetic Mountains, SE Spain (Carretero et al., 2010). This species may be found in sympatry with Podarcis hispanica ss a member of the P. hispanica spe- cies complex (Carretero, 2008b; Kaliontzopoulou et al., 2011, 2012), which also inhabits rocky habitats. Correla- tive distribution models (Rubio and Carrascal, 1994; Sil- lero et al., 2009; Carretero et al., 2010) indicate that, at large scale, the presence of A. marchi depends on moun- tains, low temperatures, high precipitation and abundant forest cover, whereas, at small scale, it is constrained by terrain roughness and closure. Although both species can be found in syntopy (Carretero et al., 2010), the general trend is that A. marchi is restricted to the more humid spots, on the contrary P. hispanica ss inhabits drier and warmer rocky habitats, therefore we can say that P. his- panica ss is more tolerant than A. marchi. To elucidate to what extent this is due to ecophysiological differences or to (mutual) exclusion, we here analyse Tp and Wl experi- mentally in both species to determine whether these traits 1) differ intrinsically between them; and 2) are affected by intraspecific and interspecific interference. MATERIALS AND METHODS Five adult males of each species were collected in a syn- topic area (Rambla los Vaquerizos, 38º3´N, 2º29´E). P. hispanica ss was assessed following Kaliounzoupoulou et al. (2011) where this samples were genetically analyzed. All individuals were captured by hand or noose (García-Muñoz and Sillero, 2010) in July 2010, and brought to the laboratory. Individuals were marked using a marker pen and kept in individual terraria for a total of eight days with food (Tenebrio molitor larvae and grass- hoppers) and water provided ad libitum and under light (14 h light – 10 h darkness) and temperature (23.3 ºC ± 1 ºC) con- ditions similar to the natural regime. Each lizard was subjected to two experiments (see below). After each set of experiments, lizards were supplied with water and food ad libitum until they recovered the initial weight, and at the end of all experiments lizards were released in the capture site. Lizards were exposed to a classic Tp experimental design of thermal gradient (~25-45 ºC, 0.3 × 0.4 × 1.0 m length experi- mental terrarium, Veríssimo and Carretero, 2009) produced by a 100 W infrared reflector bulb fixed 15 cm above the substrate and maintaining natural photoperiod. The walls of the terrari- um were covered with white panels to prevent external stimuli influencing the lizard’s thermoregulatory behaviour. Tp was measured with k-thermocouple digital thermometer (HIBOK 14; manufactured by HIBOK, precision 0.01 ºC) by inserting a probe in the cloaca every hour during 10 hours (from 9:00 to 18:00 h local time). Snout ventral length (SVL) of each lizard was also registered to the nearest 0.01 mm using a digital cal- liper (model: Mitutoyo Digimatic Caliper). Evaporative water loss experiment were developed using sealed chambers (~25 ºC controlled temperature chamber, ~30-35 %RH generated by 100 g of silica gel). Wl was meas- ured with a precision balance (precision 0.0001 mg; CPA model 224S, Sartorius Bohemia, New York). Initial weight (W0) at the start of the evaporative water loss experiment was recorded. Lizards were individually placed in transparent plastic boxes covered with a perforated lid and floor placed inside another box with 5 g of silica gel at the bottom that allowed free air exchange between the individual box and sealed chambers. Five small boxes with lizards were placed in a sealed chamber at the same time. Lizards inside the boxes were removed from the chamber hourly, weighed together with the box on the preci- sion balance and put back into the chamber, over a period of 12 hours. The whole operation did not take longer that 20 sec. The sealed chambers were maintained in obscurity and an undis- turbed room in order to avoid stressing the animals. Values of Tp (untransformed) did not deviate from nor- mality (Shapiro-Wilk’s test, P > 0.05 in all cases), were homo- scedastic (univariate Levene’s tests and multivariate Box M, P > 0.05 in all cases) and variances and means were uncorrelated. Repeat measures (rm) ANOVA, and rmANCOVAs (with LogS- VL; and with LogSVL and LogW0 (initial weight) as covari- ables) were performed in order to evaluate differences in Tp (hourly Tp; ºC) between both species (SP; A. marchi and P. his- panica s.s.) A rmANOVA, and two rmANCOVAs (with LogS- VL, and with logSVL and initial weight, LogW0 as covariables) were performed to evaluate differences on accumulative water loss (Wl = Log {[(Wo-Wn)/Wo]*100}). A post-hoc Duncan´s test was performed after rmANOVA and rmANCOVA in order to detect differences (with and without body size/mass effects) in Wl between species. In addition, Pearson correlation and linear least-squares were performed (for both species) in order to determine if there 125Comparative ecophysiology of two sympatric lizards is a relationship between Tp (Mean Tp for the 10 h, ºC) and total Wl (for the total accumulative water loss 12 h; Wl = Log (W8-W20)). RESULTS Lizards of both species showed similar preferred body temperatures (Algyroides marchi, mean ± SD Tp = 31.5 ± 0.5; P. hispanica ss, mean ± SD Tp = 31.8 ± 1.5) and repeated measures AN(C)OVA showed no statistical differences, also after size-mass correction (rmANOVA, F9, 72 = 1.702, P = 0.104; rmANCOVA, F9, 63 = 0.394, P = 0.933, LogSVL = 3.830; rmANCOVA, F9, 54 = 0.293, P = 0.973, LogSVL = 3.830, LogW0 = 0.776). In water loss experiments, A. marchi showed the highest Wl rates, reaching at the end of the experiment a water loss of approximately 10% of the initial weight. In contrast, P. hispanica ss showed much lower rates, espe- cially during the first hours, final water loss attaining 3% of the initial weight. In addition, both species showed different Wl rates though time (Time*Sp, rmANOVA, Fig. 1) and post-hoc analyses revealed that both spe- cies responded in different ways to Wl in the first hours (Duncan test: Wl1-4, P = 0.037; Wl1-5, P = 0.035). These differences between both species were not SVL- or weight dependent, and these interactions remained significant (Time*Sp; rmANCOVAs, Table 2) after size/body mass was accounted for. In addition, another post-hoc test was performed after including covariables to test if the differ- ences between both species were SVL- or weight depend- ent or intrinsic (Duncan test: Cov LogSVL, Wl1-3, P = Fig. 1. Cumulative water loss in both species tested (A. marchi; P. hispanica ss), different whiskers denote different cumulative water loss intervals and their 95% IC, Wl = Log{[(W0-Wn)/W0]*100}, for both species in an experiment conducted during 12 hours. Fig. 2. Relationship among preferred temperature (mean; Log transformed) and total water loss (Log transformed) in both species 126 Enrique García-Muñoz, Miguel Angel Carretero 0.046; Wl1-4, P = 0.021; Wl1-5, P = 0.020; Wl1-6, P = 0.029; Duncan test: CovW0, Wl1-4, P = 0.029, Wl1-5, P = 0.027, Wl1-6, P = 0.039). Regression between Tp (mean Tp 10 hours, ºC) and Wl (12 hours, total accumulative water loss) varied between species. While in A. marchi no significant rela- tions were detected, P. hispanica ss showed an inverse relationship between Tp and Wl (Fig. 2). DISCUSSION Our results indicate that both sympatric lizards dif- fer in some ecophysiological traits. Concerning the ther- mal traits, both species appeared to have similar pre- ferred temperatures which remained not significant after variation in body mass/size was accounted for. As other lacertid lizards, A. marchi and P. hispanica ss kept their body temperatures within a narrow range when free of thermal constraints (reviewed in Castilla et al., 1999; Car- retero et al., 2005; Carretero at al., 2006; Veríssimo and Carretero, 2009). It is well documented that behavioural adjustments are the primary means by which lizards buff- er fluctuations in ambient heat loads to maintain their Tp within the range that is conducive to optimal perfor- mance (Castilla et al., 1999). Regarding the water ecophysiology, much less is known in lizards in general, and only two very recent stud- ies analysed lacertid species (Carneiro et al., in press; Oso- jnik et al., 2013). Nevertheless, physiological studies have provided abundant information on osmoregulation and on mechanisms of water conservation of other lizards (Min- nich, 1982; Nagy, 1982). Thus, the resistance to water loss reflects the combined effect of two integumentary compo- nents on the rate of water loss. The first one is a structural component, including differences in skin micro-structures and lipid content. The second component is more dynam- ic, representing physiological, vasomotor responses to short-term variations in the environment. This physiologi- cal response enables better and more immediate control of Wl (Eynan and Dmi´el, 1993). Our results not only show intraspecific differences in the magnitude of absolute water loss rate but also provide evidence for a dissimilar pattern of water loss throughout a normal diel activity period. Under the same conditions, A. marchi lost more water and at a more even rate than P. hispanica ss. In fact, P. hispanica ss displayed an apparent response to dehydration within the first four hours while A. marchi did not. In addition, results for P. hispanica ss supported a trade-off between Tp and Wl at the individual level while those for A. marchi were inconclusive. Future studies should evaluate such a relationship in others species. Altogether, our ecophysiological results suggest that water should be more important than temperature to explain the distribution patterns of A. marchi vs. P. hispan- ica ss, the former risking dehydration in dry areas what- ever the temperature. This is accordance with high resolu- tion distribution models (Sillero et al., 2009; Carretero et al., 2010) indicating that A. marchi is restricted to moun- tains, high precipitation and good forest cover at a large scale and rugged and steep terrains at a small scale. These humid environments (allow to minimise lizard evaporative loss) are scarce in Mediterranean areas, which are charac- terised by midday and aestival draught, particularly in SE Iberia. Hence, it is not surprising that A. marchi displays such a restricted distribution. Also, taking into account the episodes of aridification that the whole Mediterranean Basin suffered periodically since the Pliocene (Cavazza and Wezel, 2003; Jiménez-Moreno et al., 2010), it would be expectable that A. marchi became gradually replaced by P. hispanica ss in this region. Going further, it is tempt- ing to associate the climate transition, from subtropical to Mediterranean, since the Miocene to the spread and diver- sification of Podarcis sp. (Carretero, 2008b) in contrast to the retraction of Algyroides ssp. in southern Europe (Har- ris et al., 1999). This, however, needs confirmation by fur- ther comparative studies with other species of both gen- era. Last but not least, if the scenarios for climate change (IPCC, 2007) are confirmed, the vulnerability of A. marchi to dehydration and competitive displacement by sympatric P. hispanica ss put the former species at a serious extinc- tion risk, in fact higher than suggested by correlation mod- els (Carvalho et al., 2010). Overall, ecophysiological traits like preferred temperatures and water loss rates represent a promising tool to build future mechanistic models for lac- ertids and likely for other lizard groups. ACKNOWLEDGEMENTS The Junta de Andalucía provided permits for sam- pling (SGYB/FOA/AFR). Funding for research was provid- ed by a PTDC/BIA–BEC/101256/2008 project awardfrom Fundaçao para a Ciência e a Tecnologia (FCT, Portugal). EG-M was supported by a postdoctoral grant from FCT (SFRH/ BPD/72806/2010). REFERENCES Adolph, S.C., Porter, W.P. (1993): Temperature, activity, and lizard life histories. Am. Nat. 142: 273-295. Angilletta, M.J., Niewiarowski, P.H., Navas C.A. (2002): The evolution of thermal physiology in ectotherms. J. Therm. Biol. 27: 249-268. 127Comparative ecophysiology of two sympatric lizards Angilletta, M.J. Jr. (2009): Thermal adaptation. Oxford University Press, Oxford. Arnold, E.N., Arribas, O.J., Carranza, S. (2007): Systemat- ics of the palaearctic and oriental lizard tribe lacertini (squamata: lacertidae: lacertinae), with descriptions of eight new genera. Zootaxa 1430: 1-8. Bauwens, D., Garland, T. Jr., Castilla, A.M., Van Damme, A. (1995): Evolution of sprint speed in lacertid lizards: morphological, physiological, and behavioural covari- ation. 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