Félix Benjamín Cruz

The interplay between claw morphology and microhabitat use in neotropical iguanian lizards.

Traditionally, it has been suggested that variation in locomotor mode should be correlated with variation in the anatomy of the structures responsible for locomotion. Indeed, organisms can expand their ecological niche by using specialized traits of the locomotor system including hooks, claws, adhesive pads, etc. Despite the fact that claws are the most common biological mechanism of clinging in vertebrates, little is known about their function or evolutionary relationship to habitat use. The present study focuses on claw morphology in 57 species of iguanian lizards occupying different microhabitats. Qualitative differences in claw shape were explored by means of digital photographs, and quantitative measurements of the length, height and curvature of the claws of both fingers and toes were taken and correlated to information on microhabitat use obtained from the literature. Our analyses showed a strong phylogenetic component that obscured relationships between morphology and ecology. Our results also show differences in claw morphology between species that appear to be related to microhabitat use (climbing versus terrestrial species), with the best ecological descriptors being claw length and height. Performance measures and biomechanical analyses of claw function may consequently be better suited to explain the evolution of claw shape in relation to habitat use in this group.

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.

Effects of different substrates on the sprint performance of lizards.

SUMMARY 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.

The Reproductive Cycle and the Size at Maturity of Tupinambis rufescens (Sauria: Teiidae) in the Dry Chaco of Argentina

The reproductive patterns of Tupinambis rufescens have not been described. The present study had three objectives: (1) to describe the reproductive and fat body cycles of T. rufescens from our study site in the dry chaco of Argentina; (2) to elucidate and analyze the distribution of sizes at which female T. rufescens reach sexual maturity; and (3) to discuss the conservation implications of the repro- ductive cycle of T. rufescens. Oviductal width increased significantly after females reached 320 mm SVL, and we presumed the increase was due to stretching of the oviducts in females that had been gravid. Seventy-eight percent of the females >319 mm snout-vent length (SVL) and 84.3% > 349 mm SVL possessed oviducts > 3.5 mm wide, our criterion for indicating a female had reproduced, while only 6.9% of the females 3.5 mm wide were convoluted, while all the narrower oviducts were straight or striated. The reproductive and fat body cycles of T. rufescens at our study site were characteristic of lizards from seasonal environments, and appeared closely tied to the onset of the rainy season and increasing spring temperatures. Mature females exhibited significantly longer ovaries in No- vember than in other months, and nesting was observed in November and December. The average clutch size was 21.4, and the smallest female we found with eggs was 330 mm SVL. Testis mass was significantly greater in November than in other months, and declined slightly throughout the activity season. Some gonadal development in males presumably occurred while the lizards were inactive during winter. Fat body volume was tightly linked to the gonadal cycle of both males and females. Females apparently allocated fat stores to developing eggs, while males probably used fat stores for spermatogenesis or for meeting the energetic demands of mate seeking. Large numbers of Tupinambis are exploited for their skins, and this study provides life history infor- mation needed to develop scientifically-based management plans, for example, the size distribution of reproductive females, the timing of reproduction, and clutch size. Based on our analyses, management strategies aimed at harvesting adults and subadults could be evaluated because the proportions of the harvest comprised of adults can be reliably determined. We recommend that strategies aimed at classifying harvests according to adults and subadults use 350 mm SVL as a cut-off point for mature females.

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).

Running in cold weather: morphology, thermal biology, and performance in the southernmost lizard clade in the world (Liolaemus lineomaculatus section: Liolaemini: Iguania)

The integration or coadaptation of morphological, physiological, and behavioral traits is represented by whole-organism performance traits such as locomotion or bite force. Additionally, maximum sprint speed is a good indicator of whole-organism performance capacity as variation in sprinting ability can affect survival. We studied thermal biology, morphology, and locomotor performance in a clade of Liolaemus lizards that occurs in the Patagonian steppe and plateaus, a type of habitat characterized by its harsh cold climate. Liolaemus of the lineomaculatus section display a complex mixture of conservative and flexible traits. The phylogenetically informed analyses of these ten Liolaemus species show little coevolution of their thermal traits (only preferred and optimum temperatures were correlated). With regard to performance, maximum speed was positively correlated with optimum temperature. Body size and morphology influenced locomotor performance. Hindlimbs are key for maximal speed, but forelimb length was a better predictor for sustained speed (i.e. average speed over a total distance of 1.2 m). Finally, sustained speed differed among species with different diets, with herbivores running on average faster over a long distance than omnivores.

Relationships among morphology, clinging performance and habitat use in Liolaemini lizards

The central tenet of ecomorphological theory holds that different ecological requirements lead to different organismal designs (morphology). Here, we studied the relationships between performance (interlocking grasping) and forelimb morphological traits in species of lizards that exploit different structural habitats in a phylogenetic context. The performance (measured by the maximum force of clinging to substrate) was measured on different substrate types. After phylogenetically informed analyses, we found that arboreal and saxicolous species showed stronger resistance to mechanical traction in all substrates when compared to generalists and sand dweller lizards. These species showed a positive relationship between forelimb dimensions (humerus length and length of claw of toe 5) and maximum force exerted, on the contrary, hand width, claw height (CH) of digits III and IV and claw length of toe 4 showed a negative relationship. In addition, we observed a partial positive correlation between CH and maximal cling force on rough surfaces, but not on smooth surfaces.

Refuge Use in a Patagonian Nocturnal Lizard, Homonota darwini: The Role of Temperature

Abstract The thermal quality of diurnal refuges is important to the performance and survival of nocturnal reptiles. We studied refuge use on both slopes of an east–west-oriented hill by the thigmothermic gecko Homonota darwini, the southernmost-distributed nocturnal lizard in the world, in the vicinity of Bariloche, Rio Negro, in the Patagonia of Argentina. Because of the harsh climatic conditions in Patagonia, suitable refuges are limited, and retreat-site use is important for these geckos. Homonota darwini used refuges significantly more frequently on the warmer western slope in our study site. Geckos on the western slope used those refuges with higher temperatures regardless of size and thickness of rocks that acted as retreats. We tested whether refuge temperature affected locomotor performance of these lizards. Performance experiments showed that maximum sprint speed was affected by the temperature of the refuges. Refuges at 22.5°C allowed lizards to achieve their fastest sprint performance. Unexpectedly, sprint performance of lizards that used refuges with temperatures >32°C was the lowest among all tested refuge temperatures (18°, 22.5°, 27.5°, and 33°C). Our data illustrate the importance of the thermal quality of refuges for reptiles living in extreme environments.

Predation on snakes of Argentina: Effects of coloration and ring pattern on coral and false coral snakes

The occurrence of coral snake coloration among unrelated venomous and non‐venomous snake species has often been explained in terms of warning coloration and mimicry. In Argentina, no field tests have been conducted to confirm this mimetic association between one venomous coral species (Micrurus phyrrocryptus, Elapidae) and two non‐venomous snake species with a similar color pattern (Lystrophis pulcher and Oxyrhopus rhombifer, Colubridae). The aims of this work were to test for the possible aposematic or cryptic function of the ring pattern and coloration of coral snakes and false coral snakes from central Argentina, and to analyse whether the pattern is effective throughout the year. Predation on snakes was estimated by using non‐toxic plasticine replicas of ringed venomous and non‐venomous snakes and unbanded green snakes placed along transects in their natural habitat during the dry and rainy season. Ringed color pattern was attacked by predators despite the background color. One of the replica types was attacked more than expected during the dry season, suggesting that both shape and width of rings may influence the choice by predators. The reaction of predators towards replicas that mimic snake species with ringed patterns is independent of the geographical region, and we can conclude that mimicry characteristics are quite general when the true models are present in the area.

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.