Relationship between oxidative stress and sexual coloration of lizards depends on thermal habitat.

Sexual signals serve as an honest indicator of individual quality, reflecting either developmental and/or maintenance costs. A possible underlying physiological mechanism is oxidative stress, which could mediate energy trade-offs between sexual signals and other quality traits. In ectotherms, thermal performance acts as a key indicator of individual quality and influence signal intensity. We investigated how oxidative state is reflected in visual signals of lizards from different thermal habitats. According to our hypothesis, efficient thermoregulation requires different strategies in different thermal environments. In a habitat with predictable temperature changes, animals are less exposed to suboptimal temperature ranges and selection will, therefore, be stronger on the maximum oxidative damage at optimal body temperature. Contrarily, in a habitat with rather stochastic thermal shifts, individuals are often constricted by suboptimal thermal conditions, and oxidative damage can be limiting on a wide temperature range. We used Iberolacerta cyreni and Psammodromus algirus inhabiting stochastic and predictable thermal environments respectively. We examined two aspects of oxidative stress: the level of reactive oxygen metabolites at the preferred temperature (maximal ROM) and the temperature range in which animals produce at least 80% of the maximum level of reactive oxygen metabolites (effective ROM range). In I. cyreni, we found that duller coloration was related to a wider effective ROM range, while expression of coloration in P. algirus was negatively correlated with the maximal ROM. Our results suggest that different thermal constraints affect different aspects of oxidative damage which can indicate individual quality and are, therefore, represented in sexual ornaments.

Environment-dependence of behavioural consistency in adult male European green lizards (Lacerta viridis).

Understanding the background mechanisms affecting the emergence and maintenance of consistent between-individual variation within population in single (animal personality) or across multiple (behavioural syndrome) behaviours has key importance. State-dependence theory suggests that behaviour is 'anchored' to individual state (e.g. body condition, gender, age) and behavioural consistency emerges through behavioural-state feedbacks. A number of relevant state variables are labile (e.g. body condition, physiological performance) and expected to be affected by short-term environmental change. Yet, whether short-term environmental shifts affect behavioural consistency during adulthood remains questionable. Here, by employing a full-factorial laboratory experiment, we explored if quantity of food (low vs. high) and time available for thermoregulation (3h vs. 10h per day) had an effect on activity and risk-taking of reproductive adult male European green lizards (Lacerta viridis). We focussed on different components of behavioural variation: (i) strength of behavioural consistency (repeatability for animal personality; between-individual correlation for behavioural syndrome), (ii) behavioural type (individual mean behaviour) and (iii) behavioural predictability (within-individual behavioural variation). Activity was repeatable in all treatments. Risk-taking was repeatable only in the low basking treatments. We found significant between-individual correlation only in the low food × long basking time group. The treatments did not affect behavioural type, but affected behavioural predictability. Activity predictability was higher in the short basking treatment, where it also decreased with size (≈ age). Risk-taking predictability in the short basking treatment increased with size under food limitation, but decreased when food supply was high. We conclude that short-term environmental change can alter various components of behavioural consistency. The effect could be detected in the presence/absence patterns of animal personality and behavioural syndrome and the level of individual behavioural predictability, but not in behavioural type.

Female Brazilian whiptail lizards (Cnemidophorus ocellifer) prefer males with high ultraviolet ornament reflectance.

Conspicuous colouration is an important way of social communication in many taxa. The role of ultraviolet (UV) signals in intraspecific communication has only recently been studied in lizards, and there is not a general understanding of the adaptive role of UV colouration. Colour ornaments can signal male quality in mate choice and are therefore suitable for reliably predicting the outcome of female preference. Here, we tested the potential role of UV colouration in female spatial preference in a non-territorial teiid lizard, Cnemidophorus ocellifer. We experimentally manipulated the UV reflectance of size-matched male pairs and tested the effects of our treatment on females' spatial distribution. We found that females associated with males of higher UV reflectance, suggesting that UV colour can be an important clue during mate preference decisions. Our results provide the first empirical evidence for the importance of UV colouration in female preference in a mutually ornamented lizard species.

Effects of energy and thermoregulation time on physiological state and sexual signal in a lizard.

Theory of sexual selection states that males often develop showy signals, which reduce their survival but increase their reproductive success. During mate choice, these conspicuous signals can be honest indicators of individual quality conveying information about the signaler's physiological state. Sexually selected signals are influenced by many environmental factors; however, whether signals and physiological state are affected together is rarely studied. The ultraviolet-blue throat color of male Lacerta viridis is an intra- and intersexually selected signal connected to blood parasite infection and influenced by environmental factors. The aim of this study was to experimentally investigate how ecologically relevant environmental factors affect color signal intensity and key physiological traits parallel. During the mating season, we exposed 40 adult male lizards infected with blood parasites to food and basking time treatments in a full factorial design. We measured color, amount of reactive oxygen metabolites (ROMs), hematocrit, immunocompetence, and blood parasite intensity before and after treatments. High basking time resulted in elevated immunocompetence coupled with increased ROMs. The high food treatment increased nuptial color brightness, but also increased ROMs and decreased immunocompetence. In summary, our study provides experimental evidence about environmentally induced parallel changes in an honest sexual signal and several quality-indicator physiological traits. We showed that available energy and time for high metabolism have independent and sometimes opposite effects on individual state.

European green lizard (Lacerta viridis) personalities: Linking behavioural types to ecologically relevant traits at different ontogenetic stages.

Consistent individual differences within (animal personality) and across (behavioural syndrome) behaviours became well recognized during the past decade. Nevertheless, our knowledge about the evolutionary and developmental mechanisms behind the phenomena is still incomplete. Here, we explored if risk-taking and exploration were consistent and linked to different ecologically relevant traits in wild-caught adult male European green lizards (Lacerta viridis) and in their 2-3 weeks old laboratory-reared offspring. Both adults and juveniles displayed animal personality, consistency being higher in juveniles. We found correlation between risk-taking and exploration (suggestive of a behavioural syndrome) only in adults. Juveniles were more explorative than adults. Large or ectoparasite-free adult males were more explorative than small or parasitized males. Juvenile females tended to be more risk-taking than males. Behaviour of fathers and their offspring did not correlate. We conclude that European green lizards show high behavioural consistency and age is an important determinant of its strength and links to traits likely affecting fitness.

Negative correlation between nuptial throat colour and blood parasite load in male European green lizards supports the Hamilton-Zuk hypothesis.

During female mate choice, conspicuous male sexual signals are used to infer male quality and choose the best sire for the offspring. The theory of parasite-mediated sexual selection (Hamilton-Zuk hypothesis) presumes that parasite infection can influence the elaboration of sexual signals: resistant individuals can invest more energy into signal expression and thus advertise their individual quality through signal intensity. By preferring these males, females can provide resistance genes for their offspring. Previous research showed that nuptial throat colour of male European green lizard, Lacerta viridis, plays a role in both inter- and intrasexual selections as a condition-dependent multiple signalling system. The aim of this study was to test the predictions of the Hamilton-Zuk hypothesis on male European green lizards. By blood sampling 30 adult males during the reproductive season, we found members of the Haemogregarinidae family in all but one individual (prevalence = 96%). The infection intensity showed strong negative correlation with the throat and belly colour brightness in line with the predictions of the Hamilton-Zuk hypothesis. In addition, we found other correlations between infection intensity and other fitness-related traits, suggesting that parasite load has a remarkable effect on individual fitness. This study shows that throat patch colour of the European green lizards not only is a multiple signalling system but also possibly acts as an honest sexual signal of health state in accordance with the Hamilton-Zuk hypothesis.

Temperature, but not available energy, affects the expression of a sexually selected ultraviolet (UV) colour trait in male European green lizards.

BACKGROUND: Colour signals are widely used in intraspecific communication and often linked to individual fitness. The development of some pigment-based (e.g. carotenoids) colours is often environment-dependent and costly for the signaller, however, for structural colours (e.g. ultraviolet [UV]) this topic is poorly understood, especially in terrestrial ectothermic vertebrates. METHODOLOGY/PRINCIPAL FINDINGS: In a factorial experiment, we studied how available energy and time at elevated body temperature affects the annual expression of the nuptial throat colour patch in male European green lizards (Lacerta viridis) after hibernation and before mating season. In this species, there is a female preference for males with high throat UV reflectance, and males with high UV reflectance are more likely to win fights. We found that (i) while food shortage decreased lizards' body condition, it did not affect colour development, and (ii) the available time for maintaining high body temperature affected the development of UV colour without affecting body condition or other colour traits. CONCLUSIONS/SIGNIFICANCE: Our results demonstrate that the expression of a sexually selected structural colour signal depends on the time at elevated body temperature affecting physiological performance but not on available energy gained from food per se in an ectothermic vertebrate. We suggest that the effect of high ambient temperature on UV colour in male L. viridis makes it an honest signal, because success in acquiring thermally favourable territories and/or effective behavioural thermoregulation can both be linked to individual quality.

Ultraviolet nuptial colour determines fight success in male European green lizards (Lacerta viridis).

Animal communication through colour signals is a central theme in sexual selection. Structural colours can be just as costly and honest signals as pigment-based colours. Ultraviolet (UV) is a structural colour that can be important both in intrasexual competition and mate choice. However, it is still unknown if a UV signal alone can determine the outcome of male-male fights. European green lizard (Lacerta viridis) males develop a nuptial throat coloration with a strong UV component. Among males differing only in their manipulated UV colour, females prefer males with higher UV. Here, we experimentally decreased the UV coloration of randomly chosen males from otherwise similar male pairs to test the hypothesis that a difference in UV colour alone can affect fight success during male-male competition. Our results fully supported the hypotheses: in almost 90 per cent of the contests the male with reduced UV lost the fight. Our results show that UV can be an important signal, affecting both female mate choice and determining male fight success.