Hot rocks or no hot rocks: overnight retreat availability and selection by a diurnal lizard

I used radio telemetry to determine the effects of substrate size and composition on overnight retreat site selection by western fence lizards ( Sceloporus occidentalis). In watersheds of northern California (USA), these lizards occupy two habitat types differing in substrate characteristics: rocky cobble bars found in the dry, active channels of rivers and grassy upland meadows. Rocky substrates, found almost exclusively on cobble bars, provided warmer potential retreat sites than all available retreat sites on meadows during the first 5 h of inactivity. Only cobble and sand substrates provided retreats with temperatures within the preferred daily active range (32–36°C) during the inactive period for these lizards (1900–0900 hours). Females on a cobble bar used rocks as retreats on >90% of nights during the breeding season whereas females on a meadow used wood (>70% of nights) and burrows (>25% of nights). In contrast to females, cobble bar males used rocks significantly less frequently (<70%) and slept in the open air significantly more frequently (25% vs. <1%). Cobble bar females further, showed a significant preference for cobbles 15 cm thick, whereas the rocks used by males did not differ significantly in thickness from those measured in randomly placed transects. Rocks 15 cm thick were the warmest retreats commonly available on this habitat type. Thus, thermal microenvironments available to and chosen by gravid female lizards differ considerably between river and non-river habitats.     

An analysis of the relative roles of plasticity and natural selection in the morphology and performance of a lizard (<Emphasis Type="Italic">Urosaurus ornatus</Emphasis>)

Evolutionary ecologists have devoted substantial attention to understanding which factors dictate processes of mortality within populations. Our goal was to understand the dynamics of natural selection on two performance traits (bite force and sprint speed) and associated morphological variables. We first quantified performance and morphology for a sample of marked tree lizards (Urosaurus ornatus) at the middle of the breeding season. We then sampled the same population in the nonbreeding season to determine which of the original lizards survived, and we also remeasured morphological and performance variables for surviving lizards. We found evidence for directional selection favoring fast sprinters in male lizards, but also a nonsignificant stabilizing trend that disfavored the very fastest lizards. However, we also detected substantial seasonal plasticity in bite force and head width, suggesting that an analysis of selection on only preselection (breeding season) values may be overly simplistic. Urosaurus males and females with low bite forces (and narrow heads) in the breeding season generally increased their bite forces and head widths during the nonbreeding season. In contrast, lizards that were initially strong biters in the breeding season diminished in head width and declined dramatically in bite force (up to about 35%). We suggest that seasonal plasticity could act as a retarding force for selection on performance, and could dampen seasonal and year-to-year fluctuations in selective pressures. We argue that this phenomenon may be particularly likely for performance traits important for social interactions related to breeding, such as bite force.     

Some Like It Hot: Camera Traps Unravel the Effects of Weather Conditions and Predator Presence on the Activity Levels of Two Lizards

It is generally assumed that favourable weather conditions determine the activity levels of lizards, because of their temperature-dependent behavioural performance. Inactivity, however, might have a selective advantage over activity, as it could increase survival by reducing exposure to predators. Consequently, the effects of weather conditions on the activity patterns of lizards should be strongly influenced by the presence of predators. Using remote camera traps, we test the hypothesis that predator presence and weather conditions interact to modulate daily activity levels in two sedentary cordylid lizards, Karusasaurus polyzonus and Ouroborus cataphractus. While both species are closely related and have a fully overlapping distribution, the former is a fast-moving lightly armoured lizard, whereas the latter is a slow-moving heavily armoured lizard. The significant interspecific difference in antipredator morphology and consequently differential vulnerability to aerial and terrestrial predators, allowed us to unravel the effects of predation risk and weather conditions on activity levels. Our results demonstrate that K. polyzonus is predominantly active during summer, when ambient temperatures are favourable enough to permit activity. In contrast, a peak in activity during spring was observed in O. cataphractus, with individuals being inactive during most of summer. While favourable weather conditions had a strong effect on the activity levels of K. polyzonus, no such relationship was present in O. cataphractus. Contrary to our hypothesis, the presence of terrestrial predators does not seem to affect daily activity levels or alter the influence of weather conditions on activity levels. We conclude that inactivity in O. cataphractus appears to be related to seasonal differences in vulnerability to predators, rather than the presence of predators, and highlight the importance of additional selective pressures, such as food abundance, in determining the species’ activity levels.     

Fitness effects of the timing of hatching may drive the evolution of temperature-dependent sex determination in short-lived lizards

In several species of short-lived Australian agamid lizards, an individual’s sex is determined by the nest temperatures encountered during incubation. The adaptive significance of such systems remains unclear. Here, we explore the hypothesis that (1) the optimal timing of hatching differs between the sexes, and thus (2) temperature-dependent sex determination (TSD) enhances maternal and offspring fitness by generating seasonal shifts in offspring sex ratios. Our model predicts that TSD can indeed enhance maternal fitness returns in short-lived lizards if (1) male–male competition is intense, thus reducing mating success of newly-matured males (but not females), and (2) the nesting season is prolonged, such that seasonal effects become significant. Available data on the distribution of TSD in Australian agamid lizards broadly support these predictions. Because both the level of male–male competition and the length of nesting season can vary at small spatial and temporal scales, selective forces on sex-determining mechanisms also should vary. Hence, our model predicts extensive small-scale (intraspecific) variation in sex-determining systems within agamid lizards, as well as among species.     

Cool shade and not-so-cool shade: How habitat loss may accelerate thermal stress under current and future climate

Worldwide habitat loss, land-use changes, and climate change threaten biodiversity, and we urgently need models that predict the combined impacts of these threats on organisms. Current models, however, overlook microhabitat diversity within landscapes and so do not accurately inform conservation efforts, particularly for ectotherms. Here, we built and field-parameterized a model to examine the effects of habitat loss and climate change on activity and microhabitat selection by a diurnal desert lizard. Our model predicted that lizards in rock-free areas would reduce summer activity levels (e.g. foraging, basking) and that future warming will gradually decrease summer activity in rocky areas, as even large rocks become thermally stressful. Warmer winters will enable more activity but will require bushes and small rocks as shade retreats. Hence, microhabitats that may seem unimportant today will become important under climate change. Modelling frameworks should consider the microhabitat requirements of organisms to improve conservation outcomes.     

Correlational selection on lay date and life-history traits: experimental manipulations of territory and nest site quality

Abundant evidence suggests that females may engage in mate choice to gain nongenetic (material) benefits from high-quality territories; however, the selective consequences that influence those choices are not well understood. We studied the fitness effects of territory quality and incubation temperature on juvenile lizards in nature. We manipulated territory quality by redistributing rocks between pairs of neighboring home ranges. Rock manipulations set up adjacent plots that were either experimentally improved or reduced in quality. We incubated eggs from field-caught gravid females in each of three temperature treatments in the laboratory (low, medium, and high temperature). Progeny were released in either experimentally improved or reduced-quality plots upon hatching, and the following spring we measured survival as a function of egg size and laying date. We conducted concurrent studies of the thermal environment on experimental territories. Improved territories provided significantly more hours for lizards to behaviorally thermoregulate at their preferred body temperature and also provided nest sites with incubation conditions that were closer to optimal compared with reduced-quality plots. Reduced-quality plots were significantly more variable in quality. Finally, we measured significant correlational selection between egg mass and laying date on manipulated plots in two separate years. Results indicate the influence of environmental variation on correlational selection on life-history traits.     

Exposure to solar radiation drives organismal vulnerability to climate: Evidence from an intertidal limpet

Understanding the physiological abilities of organisms to cope with heat stress is critical for predictions of species’ distributions in response to climate change. We investigated physiological responses (respiration and heart beat rate) of the ectotherm limpet Patella vulgata to heat stress events during emersion and the role of seasonal and microclimatic acclimatization for individual thermal tolerance limits. Individuals were collected from 5 microhabitats characterized by different exposure to solar radiation in the high intertidal zone of a semi-exposed rocky shore in winter and summer of 2014. Upper thermal tolerance limits (heat coma temperatures – HCTs, and heart rate Arrhenius break temperatures - ABTs) were determined for individuals from each microhabitat in both seasons under laboratory conditions. While we found a clear seasonal acclimatization, i.e., higher HCTs and ABTs in summer than in winter, we did not find evidence for microhabitat-specific responses that would suggest microclimatic acclimatization. However, operative limpet temperatures derived from in-situ temperature measurements suggest that individuals from sun exposed microhabitats have a much narrower thermal safety margins than those from less exposed surfaces or within crevices. Microhabitat specific thermal safety margins caused by high thermal heterogeneity at small spatial scales and the lack of short term acclimatization will likely shape small scale distribution patterns of intertidal species in response to the predicted increase in the frequency and intensity of heat waves.     

Foraging behavior and spatial use of a rock specialist: the southern vizcacha (Lagidium viscacia), and the exotic European hare (Lepus europaeus) in rocky outcrops of northwestern Patagonia, Argentina

The southern vizcacha (Lagidium viscacia) and the exotic European hare (Lepus europaeus) are two medium-sized herbivores that inhabit rocky outcrops in Patagonian steppe. These species overlap in diet and spatial use at medium distances from rocky outcrops in summer. We evaluated the spatial use through feces distribution in winter and determined seasonal foraging intensity in relation to the distances from rocky outcrops in order to elucidate how these herbivores use food and spatial resources in food scarcity periods. The vizcacha utilized the habitat close to rocky outcrops (<40 m) independent of season, while the hare exploited the space more widely, especially distances >40 m. However, in winter, at medium distances from rocky outcrops, there was partial spatial overlap because hares' activities were closer to rocky outcrops. Foraging intensity increased significantly in areas used by the vizcacha closer to rocky outcrops when food availability decreased, and the grasses Stipa speciosa, Poa sp., and Festuca pallescens were strongly foraged. In contrast, foraging intensity showed no changes in further distances to rocky outcrops and more use by the hare. The spatial and feeding behavior of the vizcacha, restricted to vicinity of rocky outcrops, showed high vulnerability to food availability changes. In resource scarcity situations, the spatial opportunistic behavior of the hare and the overlap in diet with the vizcacha constitutes a threat to this native herbivore. It is necessary to monitor populations of hare, since high densities could lead to food competition, impacting the small colonies of the southern vizcacha.     

Rock-dwelling lizards exhibit less sensitivity of sprint speed to increases in substrate rugosity

Effectively moving across variable substrates is important to all terrestrial animals. The effects of substrates on lizard performance have ecological ramifications including the partitioning of habitat according to sprinting ability on different surfaces. This phenomenon is known as sprint sensitivity, or the decrease in sprint speed due to change in substrate. However, sprint sensitivity has been characterized only in arboreal Anolis lizards. Our study measured sensitivity to substrate rugosity among six lizard species that occupy rocky, sandy, and/or arboreal habitats. Lizards that use rocky habitats are less sensitive to changes in substrate rugosity, followed by arboreal lizards, and then by lizards that use sandy habitats. We infer from comparative phylogenetic analysis that forelimb, chest, and tail dimensions are important external morphological features related to sensitivity to changes in substrate rugosity.     

Behavioural thermoregulation by the endangered crocodile lizard (Shinisaurus crocodilurus) in captivity

Understanding the factors that may affect behavioural thermoregulation of endangered reptiles is important for their conservation because thermoregulation determines body temperatures and in turn physiological functions of these ectotherms. Here we measured seasonal variation in operative environmental temperature (T_e), body temperature (T_b), and microhabitat use of endangered crocodile lizards (Shinisaurus crocodilurus) from a captive population, within open and shaded enclosures, to understand how they respond to thermally challenging environments. T_e was higher in open enclosures than in shaded enclosures. The T_b of lizards differed between the open and shaded enclosures in summer and autumn, but not in spring. In summer, crocodile lizards stayed in the water to avoid overheating, whereas in autumn, crocodile lizards perched on branches seeking optimal thermal environments. Crocodile lizards showed higher thermoregulatory effectiveness in open enclosures (with low thermal quality) than in shaded enclosures. Our study suggests that the crocodile lizard is capable of behavioural thermoregulation via microhabitat selection, although overall, it is not an effective thermoregulator. Therefore, maintaining diverse thermal environments in natural habitats for behavioural thermoregulation is an essential measure to conserve this endangered species both in the field and captivity.     

The impact of insularity on the thermoregulation of a Mediterranean lizard

The overall biology of ectotherms is strongly affected by the thermal quality of the environment. The particular conditions prevailing on islands have a strong effect on numerous features of animal life. In this study we compared mainland and island populations of the lizard Lacerta trilineata and hypothesized that insularity would affect the thermoregulatory strategy. Continental habitats were of lower thermal quality, experiencing more intense fluctuations and had higher values of operative temperatures. Nevertheless mainland lizards selected for higher body temperatures in the lab and showed more effective thermoregulation during summer than their island peers. Lizards achieved similar body temperatures in the field in both types of habitat, underlining the importance of predation as a potential factor to mainland lizards that failed to reach their higher thermal preferences. Both island and mainland populations of L. trilineata have been adapted to their thermal environment, supporting the labile view on the evolution of thermal physiology for this species.     

Diel and seasonal variations in the thermal biology of San Cristobal Lava Lizards (Microlophus bivittatus)

Thermal biology, and therefore energy acquisition and survival, of ectotherms can be affected by diel and seasonal patterns of environmental temperatures. Galápagos Lava Lizards live in seasonal environments that are characterized by a warm and wet period when reproductive activity is maximal, and cooler and drier period. With the use of radiotelemetric techniques to record lizard surface temperatures (T_s), we studied the thermal ecology of the San Cristóbal Lava Lizard (Microlophus bivittatus) during both the warm and cool seasons over two years. During the diel activity period and when operative temperatures exceeded T_set-min, at least on rock faces without canopy, 52% or less of the T_s observations fell within the laboratory-determined T_set range (36–40 °C). Therefore, lizards may have avoided very warm midday temperatures in shaded microhabitats and the lag times in changes in T_s values occurred as operative temperatures rose rapidly during late morning warming phase. Lizards effectively thermoregulated during a year with moderate warm season temperatures and during a cool season that was unseasonably warm. In contrast, lizards less effectively thermoregulated during the warmest and coolest years of the study. We did not detect intersexual differences in thermoregulation although males may thermoregulate less effectively than do females during the cool season although we were unable to detect significant differences using our nonparametric statistical techniques.     

Microhabitats choice in intertidal gastropods is species-, temperature- and habitat-specific

Understanding how behavioural adaptations can limit thermal stress for intertidal gastropods will be crucial for climate models. Some behavioural adaptations are already known to limit desiccation and thermal stresses as shell-lifting, shell-standing, towering, aggregation of conspecifics or habitat selection. Here we used the IRT (i.e. infrared thermography) to investigate the thermal heterogeneity of a rocky platform, with four different macrohabitats (i.e. bare rock, rock with barnacles, mussels and mussels incrusted by barnacles) over four thermally contrasted months. We investigated the body temperature of Littorina littorea and Patella vulgata found on this platform and the temperature of their microhabitat (i.e. the substratum within one body length around of each individual). We also considered the aggregation behaviour of each species and assessed the percentage of thermal microhabitat choice (i.e choice for a microhabitat with a temperature different than the surrounding substrate). We did not find any aggregation of L. littorea on the rocky platform during the four studied months. In contrast, P. vulgata were found in aggregates in all the studied periods and within each habitat, but there was no difference in body temperature between aggregated and solitary individuals. These two gastropods species were preferentially found on rock covered by barnacles in the four studied months. The presence of a thermal microhabitat choice in L. littorea and P. vulgata is habitat-dependent and also season-dependent. In June, July and November the choice was for a microhabitat with temperatures lower than the temperatures of the surrounding substrate whereas in December, individuals choose microhabitats with higher temperatures than the temperatures of their substratum. Taken together, these results suggest that gastropods species are able to explore their environment to find sustainable thermal macrohabitats and microhabitats and adapt this behaviour in function of the conditions of temperatures.     

Partitioning thermal habitat on a vertical rock,a herculean task

Species occurring in sympatry have to effectively segregate their niche in order to co-exist. In the case of ectotherms in particular, the very important parameter of thermal biology has to be taken into account. Here we investigated the thermoregulatory effectiveness (E) of two endemic Greek lizards (Hellenolacerta graeca and Podarcis peloponnesiacus) that live syntopically on a rocky cliff in the Peloponnese. We presumed that the two species would select different microhabitats, to avoid interspecific competition, and follow a similar thermoregulation pattern as they experience the same conditions. We also expected that E values for both species would differ depending on the season. Overall, we found that the two species had similar E values for each season but differentiated partial thermoregulatory attributes. Though they both occurred in the same types of microhabitat, H. graeca selected higher sites (average 99 cm above ground) than P. peloponnesiacus (average 44 cm). Also, the latter achieved higher preferred temperatures during summer and winter. Finally, the effectiveness of thermoregulation for both species varied interseasonally and received its highest values during summer, in response to the lowest thermal quality that was observed then. Similar studies stress the importance of thermal shifts for ectotherm co-existence.     

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.     

Factors affecting retreat‐site selection by coppertail skinks (Ctenotus taeniolatus) from sandstone outcrops in eastern Australia

Abstract Coppertail skinks (Ctenotus taeniolatus) are medium‐sized diurnal lizards that are widespread in eastern Australia. Field surveys on sandstone outcrops in the Sydney region showed that these animals use available rocks non‐randomly: coppertails were found under large, sun‐exposed rocks on deep soil. Choice trials in the laboratory documented selection of retreat sites that were warm, with substrate preferences shifting in a diel cycle. Lizards selected retreats with rock substrates during the day but with sandy substrates at night. Rocky retreats attain higher temperatures during daylight hours, but burrows in sand beneath rocks may provide greater protection against ingress by nocturnal predators. During fieldwork we rarely found lizards under rocks with either ants or centipedes, suggesting that coppertails may avoid these predatory invertebrates. Tongue‐flick trials showed that the lizards could discriminate among common ant species based on chemosensory cues, but apparently could not detect centipede chemical cues. In experimental trials, the lizards did not avoid retreat sites scented by either ants or centipedes. Our data thus suggest that retreat‐site selection in coppertails is driven by abiotic cues (rock size, sun exposure and substrate type) that may confer fitness benefits in terms of thermoregulation and predator avoidance, with biotic cues playing a less important role.     

Field use of maximal sprint speed by collared lizards (Crotaphytus collaris): compensation and sexual selection

To understand how selection acts on performance capacity, the ecological role of the performance trait being measured must be determined. Knowing if and when an animal uses maximal performance capacity may give insight into what specific selective pressures may be acting on performance, because individuals are expected to use close to maximal capacity only in contexts important to survival or reproductive success. Furthermore, if an ecological context is important, poor performers are expected to compensate behaviorally. To understand the relative roles of natural and sexual selection on maximal sprint speed capacity we measured maximal sprint speed of collared lizards (Crotaphytus collaris) in the laboratory and field-realized sprint speed for the same individuals in three different contexts (foraging, escaping a predator, and responding to a rival intruder). Females used closer to maximal speed while escaping predators than in the other contexts. Adult males, on the other hand, used closer to maximal speed while responding to an unfamiliar male intruder tethered within their territory. Sprint speeds during foraging attempts were far below maximal capacity for all lizards. Yearlings appeared to compensate for having lower absolute maximal capacity by using a greater percentage of their maximal capacity while foraging and escaping predators than did adults of either sex. We also found evidence for compensation within age and sex classes, where slower individuals used a greater percentage of their maximal capacity than faster individuals. However, this was true only while foraging and escaping predators and not while responding to a rival. Collared lizards appeared to choose microhabitats near refugia such that maximal speed was not necessary to escape predators. Although natural selection for predator avoidance cannot be ruled out as a selective force acting on locomotor performance in collared lizards, intrasexual selection for territory maintenance may be more important for territorial males.     

Thermal regimes during incubation do not affect mean selected temperatures of hatchling lizards (Bassiana duperreyi, Scincidae)

Incubation temperatures profoundly affect many phenotypic traits of squamate reptiles, and mean selected body temperatures of such animals also are plastic in response to environmental factors. Plausibly, then, incubation temperatures might affect hatchling thermoregulation, either via adaptation (i.e., populations that historically experience different nest conditions, also will diverge in hatchling thermoregulatory behaviour) or phenotypic plasticity (incubation temperatures directly modify hatchling behaviours). We tested this hypothesis with a montane scincid lizard (Bassiana duperreyi), using thermal-imaging methods to quantify temperatures (of both head and body) selected by hatchling lizards. The young lizards kept their heads cooler than their bodies, but mean selected temperatures did not differ among hatchlings from three populations with differing thermal regimes in natural nests, nor were they affected by thermal conditions during incubation. The conservatism of mean selected temperatures stands in strong contrast to the lability of many other phenotypic traits in response to incubation temperatures in this species.     

Nurse rocks influence forest expansion over native grassland in southern Brazil

Questions: Do small rocky outcrops favor forest plant colonization and establishment in grasslands? If so, what are the potential mechanisms involved in this process? Location: Araucaria forest and Campos grassland mosaic in southern Brazilian highlands (29°29′S, 50°12′W). Methods: We collected data on the density of forest woody species in plots located on rocky outcrops and in open fields subject to different management regimes. We evaluated the influence of outcrops on the density of forest plants ≤60‐cm tall, and the effects of other environmental variables and of site on plant density; we also used information on diaspore traits of the species to discuss the way in which plants disperse to the outcrops. Results: Rocky outcrops harbored a significantly higher density of forest plants than open fields, irrespective of site effect, and rock height was the best predictor for plant density on outcrops. Diaspores of all recorded species possess characteristics associated with dispersal by birds or mammals or by both. Conclusions: Small rocky outcrops markedly influence forest expansion by acting as perches for vertebrate dispersers and as nurse objects and safe sites for plants. Forest expansion starting in small outcrops possibly occurs as follows: perching of dispersers and increase of seed rain on rocks, promotion of better conditions of establishment for forest plants by nurse rocks, protection of plants sensitive to grazing and fire, and nucleation of forest vegetation.     

Managing rock outcrops to improve biodiversity conservation in Australian agricultural landscapes

Summary  Rocky outcrops are prominent geological features in agricultural landscapes worldwide. Reptiles are a major component of these habitats and some species are restricted to, and more abundant on, rocky outcrops than in remnant vegetation. Rock outcrops are important to reptiles because they provide resources that are often limited in the surrounding landscape (e.g. micro-gradients in climatic conditions, basking- and retreat-sites). However, there is a knowledge gap in the literature addressing the conservation value of small, rocky outcrops. Management may be necessary to reverse habitat degradation in these systems. We identify four key areas of management that need to be addressed to improve outcrop habitat values and enhance biodiversity conservation in agricultural landscapes. Specific actions involve: (i) protecting outcrops from processes that cause damage to rock microhabitat, (ii) monitoring and managing changes in vegetation structure to maintain thermally suitable environments, (iii) applying integrated pest animal control and (iv) improving matrix management to enhance inselberg function and landscape connectivity. Further research is required to evaluate the efficacy of different management regimes on outcrop biota. We hope this paper will provide the stimulus for land managers to incorporate rocky outcrops in future biodiversity conservation programmes.