Thermo-physiological changes and reproductive investment in a liolaemid lizard at the extreme of the slow-fast continuum.

Gravid female lizards often experience reduced thermal preferences and impaired locomotor performance. These changes have been attributed to the physical burden of the clutch, but some authors have suggested that they may be due to physiological adjustments. We compared the thermal biology and locomotor performance of the lizard Liolaemus wiegmannii 1 week before and 1 week after oviposition. We found that gravid females had a thermal preference 1°C lower than that of non-gravid females. This was accompanied by a change in the thermal dependence of maximum running speed. The thermal optimum for locomotor performance was 2.6°C lower before oviposition than after. At relatively low temperatures (22 and 26°C), running speeds of females before oviposition were up to 31% higher than for females after oviposition. However, at temperatures above 26°C, females achieved similar maximum running speeds (∼1.5 m s-1) regardless of reproductive stage. The magnitude of the changes in thermal parameters and locomotor performance of L. wiegmannii females was independent of relative clutch mass (clutches weighed up to 89% of post-oviposition body mass). This suggests that the changes are not simply due to the clutch mass, but are also due to physiological adjustments. Liolaemus wiegmannii females simultaneously adjusted their own physiology in a short period in order to improve locomotor performance and allocated energy for embryonic development during late gravid stage. Our findings have implications for understanding the mechanisms underlying life histories of lizards on the fast extreme of the slow-fast continuum, where physiological exhaustion could play an important role.

Chasing the Patagonian sun: comparative thermal biology of Liolaemus lizards.

The importance of the thermal environment for ectotherms and its relationship with thermal physiology and ecology is widely recognized. Several models have been proposed to explain the evolution of the thermal biology of ectotherms, but experimental studies have provided mixed support. Lizards from the Liolaemus goetschi group can be found along a wide latitudinal range across Argentina. The group is monophyletic and widely distributed, and therefore provides excellent opportunities to study the evolution of thermal biology. We studied thermal variables of 13 species of the L. goetschi group, in order to answer three questions. First, are aspects of the thermal biology of the L. goetschi group modelled by the environment or are they evolutionarily conservative? Second, have thermal characteristics of these animals co-evolved? And third, how do the patterns of co-evolution observed within the L. goetschi group compare to those in a taxonomically wider selection of species of Liolaemus? We collected data on 13 focal species and used species information of Liolaemus lizards available in the literature and additional data obtained by the authors. We tackled these questions using both conventional and phylogenetically based analyses. Our results show that lizards from the L. goetschi group and the genus Liolaemus in general vary in critical thermal minimum in relation to mean air temperature, and particularly the L. goetschi group shows that air temperature is associated with critical thermal range, as well as with body temperature. Although the effect of phylogeny cannot be ignored, our results indicate that these thermal biology aspects are modelled by cold environments of Patagonia, while other aspects (preferred body temperature and critical thermal maximum) are more conservative. We found evidence of co-evolutionary patterns between critical thermal minimum and preferred body temperature at both phylogenetic scales (the L. goetschi group and the extended sample of 68 Liolaemus species).

Effects of different substrates on the sprint performance of lizards.

The variation in substrate structure is one of the most important determinants of the locomotor abilities of lizards. Lizards are found across a range of habitats, from large rocks to loose sand, each of them with conflicting mechanical demands on locomotion. We examined the relationships among sprint speed, morphology and different types of substrate surfaces in species of lizards that exploit different structural habitats (arboreal, saxicolous, terrestrial and arenicolous) in a phylogenetic context. Our main goals were to assess which processes drive variability in morphology (i.e. phylogeny or adaptation to habitat) in order to understand how substrate structure affects sprint speed in species occupying different habitats and to determine the relationship between morphology and performance. Liolaemini lizards show that most morphological traits are constrained by phylogeny, particularly toe 3, the femur and foot. All ecological groups showed significant differences on rocky surfaces. Surprisingly, no ecological group performed better on the surface resembling its own habitat. Moreover, all groups exhibited significant differences in sprint speed among the three different types of experimental substrates and showed the best performance on sand, with the exception of the arboreal group. Despite the fact that species use different types of habitats, the highly conservative morphology of Liolaemini species and the similar levels of performance on different types of substrates suggest that they confer to the 'jack of all trades and master of none' principle.

Influence of temperature variation on incubation period, hatching success, sex ratio, and phenotypes in Caiman latirostris.

Temperature is crucial for reptiles, also during embryonic development, particularly for species with temperature-dependent sex determination. Under natural conditions, Broad-snouted caiman (Caiman latirostris) eggs are influenced by thermal changes in the interior of the nest related to the external environmental temperature. As nests are subject to variations in temperature and most lab studies on crocodilian incubation have been carried out at constant temperatures, we were interested in determining how temperature fluctuations may affect the development of caiman embryos. We investigated the effects of incubation at constant temperatures (31°C, 32°C, and 33°C) and fluctuating temperatures (31 ± 2, 32 ± 1, and 32 ± 2°C) on the following aspects: incubation period duration, hatching success, sex ratio, total length, and body mass of C. latirostris hatchlings. Eggs incubated at 31°C produced 100% females, those at 32°C produced 71.6% females (however, the sex ratio was nest related), and at 33°C produced 100% males. We found a masculinizing effect when incubation was at 31 ± 2°C compared with a constant 31°C; and temperature fluctuations at 32°C (32 ± 1 and 32 ± 2°C) had a negative effect on hatchlings size and mass, and hatching success compared with constant incubation temperatures of 32°C and 33°C. Finally, the effect of temperature variation during the incubation period on sex ratio, hatching success, and phenotype depends on the mean temperature, as the fluctuation around 31°C affected the sex ratios and incubation period, and the fluctuation around 32°C affected hatchling success and size.

Anuran forelimb muscle tendinous structures and their relationship with locomotor modes and habitat use.

The interaction between organisms and their environment is central in functional morphology. Differences in habitat usage may imply divergent morphology of locomotor systems; thus, detecting which morphological traits are conservative across lineages and which ones vary under environmental pressure is important in evolutionary studies. We studied internal and external morphology in 28 species of Neotropical anurans. Our aim was to determine if internal morphology (muscle and tendons) shows lower phylogenetic signal than external morphology. In addition, we wanted to know if morphology varies in relation to the habitat use and if there are different functional groups. We found differences in the degree of phylogenetic signal on the groups of traits. Interestingly, postaxial regions of the forelimb are evolutionarily more labile than the preaxial regions. Phylomorphospace plots show that arboreal (jumpers and graspers) and swimmer frogs cluster based on length of fingers and the lack of sesamoid, also reflected by the use of habitat. These functional clusters are also related to phylogeny. Sesamoid and flexor plate dimensions together with digit tendons showed to be important to discriminate functional groups as well as use of habitat classification. Our results allow us to identify a "grasping syndrome" in the hand of these frogs, where palmar sesamoid and flexor plate are absent and a third metacarpal with a bony knob are typical. Thus, a lighter skeleton, long fingers and a prensile hand may be key for arboreality.

Are the number and size of scales in Liolaemus lizards driven by climate?

Ectothermic vertebrates are sensitive to thermal fluctuations in the environments where they occur. To buffer these fluctuations, ectotherms use different strategies, including the integument, which is a barrier that minimizes temperature exchange between the inner body and the surrounding air. In lizards, this barrier is constituted by keratinized scales of variable size, shape and texture, and its main function is protection, water loss avoidance and thermoregulation. The size of scales in lizards has been proposed to vary in relation to climatic gradients; however, it has also been observed that in some groups of Iguanian lizards it could be related to phylogeny. Thus, here, we studied the area and number of scales (dorsal and ventral) of 60 species of Liolaemus lizards distributed in a broad latitudinal and altitudinal gradient to determine the nature of the variation of the scales with climate, and found that the number and size of scales are related to climatic variables, such as temperature and geographical variables as altitude. The evolutionary process that best explained how these morphological variables evolved was the Ornstein-Uhlenbeck model. The number of scales seemed to be related to common ancestry, whereas dorsal and ventral scale areas seemed to vary as a consequence of ecological traits. In fact, the ventral area is less exposed to climate conditions such as ultraviolet radiation or wind and is, thus, under less pressure to change in response to alterations in external conditions. It is possible that scale ornamentation, such as keels and granulosity, may bring some more information in this regard.

When a general morphology allows many habitat uses.

During the last decades the study of functional morphology received more attention incorporating more detailed data corresponding to the internal anatomy that together contribute for a better understanding of the functional basis in locomotion. Here we focus on 2 lizard families, Tropiduridae and Liolaemidae, and use information related to muscle-tendinous and external morphology traits of hind legs. We investigate whether the value of the traits analyzed tend to exhibit a reduced phenotypic variation produced by stabilizing selection, and whether species showing specialization in their habitat use will also exhibit special morphological features related to it. As a result, we identified that evolution of hind limb traits is mainly explained by the Ornstein-Uhlenbeck model, suggesting stabilizing selection. Liolaemids and tropidurids show clear ecomorphological trends in the variables considered, with sand lizards presenting the most specialized morphological traits. Some ecomorphological trends differ between the 2 lineages, and traits of internal morphology tend to be more flexible than those of external morphology, restricting the ability to identify ecomorphs shared between these 2 lineages. Conservative traits of external morphology likely explain such restriction, as ecomorphs have been historically defined in other lizard clades based on variation of external morphology.

Thermal sensitivity of cold climate lizards and the importance of distributional ranges.

One of the fundamental goals in macroecology is to understand the relationship among species' geographic ranges, ecophysiology, and climate; however, the mechanisms underlying the distributional geographic patterns observed remain unknown for most organisms. In the case of ectotherms this is particularly important because the knowledge of these interactions may provide a robust framework for predicting the potential consequences of climate change in these organisms. Here we studied the relationship of thermal sensitivity and thermal tolerance in Patagonian lizards and their geographic ranges, proposing that species with wider distributions have broader plasticity and thermal tolerance. We predicted that lizard thermal physiology is related to the thermal characteristics of the environment. We also explored the presence of trade-offs of some thermal traits and evaluated the potential effects of a predicted scenario of climate change for these species. We examined sixteen species of Liolaemini lizards from Patagonia representing species with different geographic range sizes. We obtained thermal tolerance data and performance curves for each species in laboratory trials. We found evidence supporting the idea that higher physiological plasticity allows species to achieve broader distribution ranges compared to species with restricted distributions. We also found a trade-off between broad levels of plasticity and higher optimum temperatures of performance. Finally, results from contrasting performance curves against the highest environmental temperatures that lizards may face in a future scenario (year 2080) suggest that the activity of species occurring at high latitudes may be unaffected by predicted climatic changes.

Anatomy of the crus and pes of neotropical iguanian lizards in relation to habitat use and digitally based grasping capabilities.

Ecomorphological studies of lizards have explored the role of various morphological traits and how these may be associated with, among other things, habitat use. We present an analysis of selected traits of internal morphology of the hind limbs of Neotropical iguanian lizards and their relationship to habitat use. Considering that one of the most widely-held hypotheses relating to the origin of grasping is associated with the exploitation of the narrow-branch arboreal habitat, we include subdivisions of this designation as two of our ecologically defined categories of habitat exploitation for analysis, and compare lizards assigned to these categories to the features displayed by terrestrial lizards. The influence of phylogeny in shaping the morphology of lizards was assessed by using the comparative method. K values were significant for several osteological traits. Most of the K values for the variables based upon muscle and tendon morphometric characters (13 out 21), by contrast, had values <1, suggesting that their variation cannot be explained by phylogeny alone. Results of our phylogenetic and conventional ANCOVA analyses reveal that the characters highlighted through the application of the comparative method are not absolutely related to habitat in terms of the categories considered here. It appears that the bauplan of the lizard pes incorporates a morphological configuration that is sufficiently versatile to enable exploitation of almost all of the available habitats. As unexpected as conservation of internal gross morphology appears, it represents a means of accommodating to environmental challenges by apparently permitting adequacy for all situations examined.

Energetics in Liolaemini lizards: implications of a small body size and ecological conservatism.

Liolaemini lizards occur in southern South America in a variety of dietary habits across a broad latitudinal and altitudinal distribution. We studied standard metabolic rates of 19 Liolaemini species and analyzed these data using both conventional and phylogenetically informed statistics. Oxygen consumption showed a significant and positive relationship with body mass (SMR = 0.109 × body mass(0.876±0.023)), with a higher slope than that expected on the basis of the three-quarter power law model. After phylogenetically informed and conventional analyses, no significant differences in metabolic rates were found to be related to diet or elevation. We hypothesize that small body size, ecological conservatism and physiological compensation may explain the lack of differences in metabolic rates observed among these lizards.

Thermal biology of Phymaturus lizards: evolutionary constraints or lack of environmental variation?

Several aspects of the biology of Phymaturus lizards including their herbivorous diet, specialized microhabitat use, and viviparous reproductive mode are highly conserved within the group. Here, we explore two aspects of Phymaturus thermal biology and test for the co-evolution among aspects of the thermal biology in these lizards, such as thermal preferenda and critical temperatures. Secondly, we explore correlations among variation in thermal biology with elevation and latitude. To do so, we used phylogenetically based comparative analyses (PCM) together with conventional statistics. Our results show that thermal biology for Phymaturus is conservative and our data do not suggest the co-evolution of thermal variables. Moreover, we detected low levels of variation in the thermal parameters studied, and no clear relationships between climatic and thermal variables. As a significant association between climatic and thermal variables could be demonstrated for a set of syntopic Liolaemus lizards, we suggest that thermal biology in Phymaturus lizards may be evolutionarily or ecologically constrained.