The highest kingdom of Anolis: Thermal biology of the Andean lizard Anolis heterodermus (Squamata: Dactyloidae) over an elevational gradient in the Eastern Cordillera of Colombia

Vertebrate ectotherms may deal with changes of environmental temperatures by behavioral and/or physiological mechanisms. Reptiles inhabiting tropical highlands face extreme fluctuating daily temperatures, and extreme values and intervals of fluctuations vary with altitude. Anolis heterodermus occurs between 1800 m to 3750 m elevation in the tropical Andes, and is the Anolis species found at the highest altitude known. We evaluated which strategies populations from elevations of 2200 m, 2650 m and 3400 m use to cope with environmental temperatures. We measured body, preferred, critical maximum and minimum temperatures, and sprint speed at different body temperatures of individuals, as well as operative temperatures. Anolis heterodermus exhibits behavioral adjustments in response to changes in environmental temperatures across altitudes. Likewise, physiological traits exhibit intrapopulation variations, but they are similar among populations, tended to the “static” side of the evolution of thermal traits spectrum. The thermoregulatory behavioral strategy in this species is extremely plastic, and lizards adjust even to fluctuating environmental conditions from day to day. Unlike other Anolis species, at low thermal quality of the habitat, lizards are thermoconformers, particularly at the highest altitudes, where cloudy days can intensify this strategy even more. Our study reveals that the pattern of strategies for dealing with thermal ambient variations and their relation to extinction risks in the tropics that are caused by global warming is perhaps more complex for lizards than previously thought.     

The effects of thermal biology and refuge availability on the restricted distribution of an alpine lizard

Aim In an effort to disentangle the ecological processes that confine ectotherms to alpine environments, we studied the thermoregulatory and microhabitat selection behaviours of the rock lizard Iberolacerta cyreni, which is endemic to some mountains of central Spain, and of the wall lizard Podarcis muralis, which is a potential competitor of rock lizards. Location We chose three areas in the Sierra de Guadarrama (central Spain) that differed in their thermal quality [mean deviation of environmental operative temperatures from the lizards’ preferred thermal range (PTR)] and refuge availability: a pine forest (1770 m a.s.l.) in which P. muralis was the only species found, and two mixed shrub and rock sites (1770 and 1900 m a.s.l.) where both species were present. Methods In the field we collected data on refuge availability, sun exposure, body temperature (T_b) and operative temperature (T_e). Thus, we estimated the thermal habitat quality of the areas sampled and the thermoregulation accuracy and effectiveness of both species. Results The pine forest had the lowest thermal quality and refuge availability. The lower‐elevation shrub site offered the best thermal quality, but refuges were much scarcer than at the higher‐elevation site. Both species thermoregulated accurately, because mean deviations of body temperature (T_b) from PTR were considerably smaller than those of T_e. Podarcis muralis had higher T_b values than did I. cyreni, which had similar T_b values at both shrub sites, whereas P. muralis had lower T_b values at higher elevation. Overall, the thermoregulatory effectiveness (extent to which T_b values are closer to the PTR than are T_e values) of both species was similar, but whereas I. cyreni thermoregulated more efficiently at higher elevation, the opposite was true for P. muralis. At the lower‐elevation shrub site, I. cyreni remained closer to refuges than did P. muralis. Main conclusions Our results suggest that the pine forest belt might prevent the expansion of rock lizards towards lower elevations as a result of its low thermal quality and scarcity of refuges, that the thermoregulatory effectiveness of rock lizards in alpine environments depends more on refuge availability than on thermal habitat quality, and that competition with wall lizards is unlikely to explain either the distribution or the thermoregulatory effectiveness of rock lizards.     

The peak of thermoregulation effectiveness: Thermal biology of the Pyrenean rock lizard,Iberolacerta bonnali (Squamata,Lacertidae)

We studied, at 2200 m altitude, the thermal biology of the Pyrenean rock lizard, Iberolacerta bonnali, in the glacial cirque of Cotatuero (National Park of Ordesa, Huesca, Spain). The preferred thermal range (PTR) of I. bonnali indicates that it is a cold-adapted ectotherm with a narrow PTR (29.20–32.77 °C). However, its PTR (3.57 °C) is twice as wide as other Iberolacerta lizards, which may be explained by its broader historical distribution. The studied area is formed by a mosaic of microhabitats which offer different operative temperatures, so that lizards have, throughout their entire daily period of activity, the opportunity to choose the most thermally suitable substrates. I. bonnali achieves an effectiveness of thermoregulation of 0.95, which makes it the highest value found to date among the Lacertidae, and one of the highest among lizards. Their relatively wide distribution, their wider PTR, and their excellent ability of thermoregulation, would make I. bonnali lizards less vulnerable to climate change than other species of Iberolacerta. Thanks to its difficult access, the studied area is not visited by a large number of tourists, as are other areas of the National Park. Thus, it is a key area for the conservation of the Pyrenean rock lizard. By shuttling between suitable microhabitats, lizards achieve suitable body temperatures during all day. However, such thermally suitable microhabitats should vary in other traits than thermal quality, such as prey availability or predation risk. Hence, it seems that these not-thermal traits are not constraining habitat selection and thermoregulation in this population. Therefore, future research in this population may study the causes that would lead lizards to prioritize thermoregulation to such extent in this population.     

The visual perceptual range of a lizard, <Emphasis Type="Italic">Tiliqua rugosa</Emphasis>

The fragmentation of landscapes produces habitat gaps where the distance between visual landmarks may exceed the perceptual range of a species and impose navigational constraints. We estimated the visual perceptual range of the Australian sleepy lizard, Tiliqua rugosa, by releasing individuals in the centre of a cleared arena in high temperature conditions, with a 0.5-m-high bush placed either 10, 20 or 30 m from the release site. Lizards were more likely to locate those bushes and shelter under them when they were closer, and no lizards found a bush at 30 m. In addition, lizards were less likely to move from the release point when bushes were at 30 m than when they were at the two closer distances. These data suggest that for sleepy lizards the perceptual range for a 0.5-m-high bush is about 20 m. In the uncleared chenopod shrub-land where these lizards live, suitable shelter bushes are an average of 10.5 m from any point in their home range, well within their perceptual range.     

Plasticity of haemoglobin concentration and thermoregulation in a mountain lizard

The plastic capability of species to cope with the new conditions created by climate change is poorly understood. This is particularly relevant for organisms restricted to high elevations because they are adapted to cold temperatures and low oxygen availability. Therefore, evaluating trait plasticity of mountain specialists is fundamental to understand their vulnerability to environmental change. We transplanted mountain lizards, Iberolacerta cyreni, 800 m downhill to evaluate the plastic response in body condition, thermoregulation traits, haemoglobin level, and haemoparasite load. Initial measurements of body mass, total haemoglobin concentration ([Hb]), hematic parasite intensities, dorsal luminance, and thermoregulatory behaviour were resampled after two and four weeks of acclimation. We also tested whether an anti-parasitic drug reduced haemoparasite intensity. After only two weeks of acclimation to a lower elevation, lizards decreased 42% in [Hb], had 17% less parasite intensities, increased body condition by 25%, and raised by ~3% their mean preferred temperatures and their voluntary thermal maximum. The anti-parasitic treatment had no significant effect on the intensity of hematic parasites, but our results suggest that negative effects of haemoparasites on [Hb] are relaxed at lower elevation. The rapid plastic changes observed in thermal preferences, body condition, [Hb], and parasite intensity of I. cyreni demonstrate a potential plastic response of a mountain specialist. This may be adaptive under the climatic extremes typical of mountain habitats. However, there is uncertainty in whether the observed plasticity can also help overcome long term environmental changes.     

Refuge sites used by the scincid lizard Tiliqua rugosa

Abstract We examined microhabitat use by the Australian scincid lizard Tiliqua rugosa (the sleepy lizard) in chenopod shrubland. Thirty radiotagged lizards were followed during their spring period of activity (September–November 2000). Characteristics of refuges used by the lizards were compared with the perennial woody plants in 10 10 m × 10 m randomly located quadrats at each of the three sites in the study area. We found that sleepy lizards used multiple refuge sites. Perennial woody bushes constituted an important habitat component for both active and inactive lizards. Use of refuge sites by the lizards was non‐random. They used large bushes with foliage in contact with the ground (‘dome shaped’), and they sheltered under thorny bushes more frequently than expected if choice was random. Bushes that were used repeatedly tended to be larger and were more likely to be dome‐shaped. Under these large and dome‐shaped bushes, daytime temperatures were lower than under smaller bushes. Refuge use was modified as the season progressed, with lizards using larger bushes and a greater proportion of dome‐shaped bushes later in the season. We suggest that sleepy lizards are modifying their refuge site use as ambient temperatures increase, based on microclimatic needs, implying an ability to discriminate among bushes.     

Geographic variation in the life history of the sagebrush lizard: the role of thermal constraints on activity

Thermal constraints on the time available for activity have been proposed as a proximate mechanism to explain variation in suites of life history traits. The longer that an ectotherm can maintain activity, the more time it has to forage and the greater chance that it will encounter a predator and be eaten. Thus, the thermal environment may produce a trade off between growth and survival when variation in the environment favors increased activity. I used mark-recapture data from a demographic study of three natural populations of the sagebrush lizard (Sceloporus graciosus) and estimates of thermal opportunity for each population to evaluate whether variation in the thermal environment can explain patterns of growth and survival that occur over an elevational gradient. Lizards from the highest elevation population exhibited higher individual growth rates than those of lizards from lower elevation, while mortality rates increased with elevation for these populations. The covariation of fast growth and high mortality with increased thermal opportunity is the opposite trend expected if the thermal environment alone is to explain patterns of life history in these lizards. Additional factors including thermal heterogeneity in the distribution of microhabitats of lizards, adaptation to local environmental conditions, and a potential trade-off between resource acquisition and predation risk need to be addressed to obtain a satisfactory explanation of the causative mechanisms producing life history variation.     

Ecophysiology of a lacertid community in the high Moroccan mountains suggests conservation guidelines

Lizard species may differ in their ecophysiology due to adaptation, plasticity and/or phylogeny. In restrictive environments, ecophysiological differences of species living in sympatry are expected to reveal long-term evolutionary responses to the abiotic environment while competitive interactions should be limited. These influences can be disentangled by combining field monitoring with experimental tests. Here, three lacertid lizard species, Atlantolacerta andreanskyi, Scelarcis perspicillata and Podarcis vaucheri sharing high mountain habitats in Oukaimeden (High Atlas, Morocco), were studied. In the field, spatiotemporal variation of the thermal and hydric environment used by the lizards was monitored using data-loggers. In the lab, thermal and hydric ecophysiology was estimated through assessments of preferred temperatures (Tp) and water loss (WL) rates. Species differed in microhabitat use and, hence, in their exposure to variations in temperature and humidity. However, they only differed in their WL (A. andreanskyi > S. perspicillata > P. vaucheri) while their Tp were similar. Such partial differences of species in in the fundamental niche, likely derived from their long-term independent phylogenetic trajectories, can be used to predict their responses to climate and habitat shifts in this and other parts of their respective ranges. Results also confirm previous suggestions that, together with thermal physiology, hydric physiology plays a prominent role in the organisation of lizard communities in the temperate region.     

Proximate causes of altitudinal differences in body size in an agamid lizard

Body size is directly linked to key life history traits such as growth, fecundity, and survivorship. Identifying the causes of body size variation is a critical task in ecological and evolutionary research. Body size variation along altitudinal gradients has received considerable attention; however, the underlying mechanisms are poorly understood. Here, we compared the growth rate and age structure of toad‐headed lizards (Phrynocephalus vlangalii) from two populations found at different elevations in the Qinghai‐Tibetan Plateau. We used mark‐recapture and skeletochronological analysis to identify the potential proximate causes of altitudinal variation in body size. Lizards from the high‐elevation site had higher growth rates and attained slightly larger adult body sizes than lizards from the low‐elevation site. However, newborns produced by high‐elevation females were smaller than those by low‐elevation females. Von Bertalanffy growth estimates predicted high‐elevation individuals would reach sexual maturity at an earlier age and have a lower mean age than low‐elevation individuals. Relatively lower mean age for the high‐elevation population was confirmed using the skeletochronological analysis. These results support the prediction that a larger adult body size of high‐elevation P. vlangalii results from higher growth rates, associated with higher resource availability.     

Effects of hypoxia on the thermal physiology of a high-elevation lizard: implications for upslope-shifting species

Montane reptiles are predicted to move to higher elevations in response to climate warming. However, whether upwards-shifting reptiles will be physiologically constrained by hypoxia at higher elevations remains unknown. We investigated the effects of hypoxic conditions on preferred body temperatures (T_pref) and thermal tolerance capacity of a montane lizard (Phrynocephalus vlangalii) from two populations on the Qinghai–Tibet Plateau. Lizards from 2600 m a.s.l. were exposed to O₂ levels mimicking those at 2600 m (control) and 3600 m (hypoxia treatment). Lizards from 3600 m a.s.l. were exposed to O₂ levels mimicking those at 3600 m (control) and 4600 m (hypoxia treatment). The T_pref did not differ between the control and hypoxia treatments in lizards from 2600 m. However, lizards from 3600 m selected lower body temperatures when exposed to the hypoxia treatment mimicking the O₂ level at 4600 m. Additionally, the hypoxia treatment induced lower critical thermal minimum (CT_min) in lizards from both populations, but did not affect the critical thermal maximum (CT_max) in either population. Our results imply that upwards-shifting reptiles may be constrained by hypoxia if a decrease in T_pref reduces thermally dependent fitness traits, despite no observed effect on their heat tolerance.     

The temperature sensitivity along elevational gradients is more stable in maximum latewood density than tree-ring width

Tree ring-based temperature reconstructions are preferably derived from maximum latewood density (MXD) compared to tree-ring width (TRW). Although temperature signals in MXD are less dependent on site ecology, systematic analyses of the effects of elevation and slope aspect on ring formation are still lacking. Here, we assess the climate sensitivity of MXD and TRW chronologies from six larch (Larix decidua Mill.) sites across the Simplon valley in the southwestern Swiss Alps, representing elevations from 1400 to 2150 m asl on both north- and south-facing slopes. We find decreasing temperature signals with decreasing elevation in MXD and TRW, though correlation coefficients are generally higher for MXD and on the warmer and dryer south exposed slopes. While the greatest temperature signals are found for MJJA at highest elevations with r = 0.71 for MXD and r = 0.57 for TRW (both p < 0.05 and for the 1928–2009 common period), MXD still correlates significantly positive at the lowest elevation site that is ~750 m below the treeline. Our findings indicate the suitability of MXD over TRW for temperature reconstructions when using historical wood sources of unknown origin.     

Living in sympatry: The effect of habitat partitioning on the thermoregulation of three Mediterranean lizards

The ability for effective, accurate and precise thermoregulation is of paramount importance for ectotherms. Sympatric lizards often partition their niche and select different microhabitats. These microhabitats, however, usually differ in their thermal conditions and lizards have to adapt their thermoregulation behavior accordingly. Here, we evaluated the impact of habitat partitioning on the thermal biology of three syntopic, congeneric lacertids (Podarcis peloponnesiacus, P. tauricus and P. muralis) from central Peloponnese, Greece. We assessed thermoregulation effectiveness (E) using the three standard thermal parameters: body (T_b), operative (T_e) and preferred (T_pref) temperatures. We hypothesized that the microhabitats used by each species would differ in thermal quality. We also predicted that all species would effectively thermoregulate, as they inhabit a thermally challenging mountain habitat. As expected, the partition of the habitat had an effect on the thermoregulation of lizards since microhabitats had different thermal qualities. All three species were effective and accurate thermoregulators but one of them achieved smaller E values as a result of the lower T_b in the field. This discrepancy could be attributed to the cooler (but more benign) thermal microhabitats that this species occupies.     

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.     

Different trends and divergent responses to climate factors in the radial growth of Abies georgei along elevations in the central Hengduan Mountains

Climate change has been unprecedented in the last half-century. Tree growth dynamics and responses to climate warming at different elevations vary by study area due to regional diversity in site-specific climatic conditions in the central Hengduan Mountains. A. georgei is the dominant species in high-elevation montane forests in the central Hengduan Mountains. To study the response of A. georgei radial growth to climate and identify tree growth trends at different elevations, tree-ring width chronologies at four elevations across the subalpine A. georgei forest belt were built and growth-climate relationships were analyzed. The primary findings of this study were as follows: (1) radial growth rates of A. georgei decreased with elevation; (2) warming alleviated the limitation of low temperatures and abundant precipitation on tree radial growth at the highest sampling site; and (3) unlike at other elevations, the trend of trees basal area increment (BAI) at the lowest sampling site showed a significant decline over the past 20 years. This suggests the presence of an elevational inflection point, likely between 3800 m and 4000 m, where tree growth trends diverge. These results confirmed that A. georgei at higher elevation in the central Hengduan Mountains currently benefits from higher temperatures. However, the effects of drought on A. georgei at lower elevations would cause radial growth to decrease with climate warming. Therefore, it is critical to establish effective management strategies based on how A. georgei responds to climate change at various elevations.     

Bergmann's Rule rules body size in an ectotherm: heat conservation in a lizard along a 2200‐metre elevational gradient

Bergmann's Rule predicts larger body sizes in colder habitats, increasing organisms' ability to conserve heat. Originally formulated for endotherms, it is controversial whether Bergmann's Rule may be applicable to ectotherms, given that larger ectotherms show diminished capacity for heating up. We predict that Bergmann's Rule will be applicable to ectotherms when the benefits of a higher conservation of heat due to a larger body size overcompensate for decreased capacity to heating up. We test this hypothesis in the lizard Psammodromus algirus, which shows increased body size with elevation in Sierra Nevada (SE Spain). We measured heating and cooling rates of lizards from different elevations (from 300 to 2500 m above sea level) under controlled conditions. We found no significant differences in the heating rate along an elevational gradient. However, the cooling rate diminished with elevation and body size: highland lizards, with larger masses, have a higher thermal inertia for cooling, which allows them to maintain heat for more time and keep a high body temperature despite the lower thermal availability. Consequently, the net gaining of heat increased with elevation and body size. This study highlights that the heat conservation mechanism for explaining Bergmann's Rule works and is applicable to ectotherms, depending on the thermal benefits and costs associated with larger body sizes.     

Geographic variation and acclimation effects on thermoregulation behavior in the widespread lizard Liolaemus pictus

Populations at the warm range margins of the species distribution may be at the greatest risks of extinction from global warming unless they can tolerate extreme environmental conditions. Yet, some studies suggest that the thermal behavior of some lizard species is evolutionarily rigid. During two successive years, we compared the thermal biology of two populations of Liolaemus pictus living at the northern (warmer) and one population living at the southern (colder) range limits, thus spanning an 800 km latitudinal distance. Populations at the two range margins belong to two deeply divergent evolutionary clades. We quantified field body temperatures (T_b), laboratory preferred body temperatures (PBT), and used operative temperature data (T_e) to calculate the effectiveness of thermoregulation (E). During one year in all populations, we further exposed half of the lizards to a cold or a hot acclimation treatment to test for plasticity in the thermal behavior. The environment at the southern range limit was characterized by cooler weather and lower T_e. Despite that, females had higher T_b and both males and females had higher PBT in the southernmost population (or clade) than in the northernmost populations. Acclimation to cold conditions led to higher PBT in all populations suggesting that plastic responses to thermal conditions, instead of evolutionary history, may contribute to geographic variation. Lizards regulated moderately well their body temperature (E≈0.7): they avoided warm microhabitats in the northern range but capitalized on warm microhabitats in the southern range. We review literature data to show that Liolaemus species increase their thermoregulation efficiency in thermally challenging environments. Altogether, this indicates that habitats of low thermal quality generally select against thermoconformity in these lizards.     

Establishing climatic constraints shaping the distribution of alien plant species along the elevation gradient in the Alps

In this work, we analyse the role of climatic constraints in shaping the distribution of alien plant species along the elevation gradient in the European Alps. Alien species occurrence was recorded in 278 plots located beside rivers, from 100 to 2,100 m a.s.l. Climate variables were calculated from the data recorded by 145 meteorological stations and interpolated by a multiple regression approach. Both richness and occurrence of aliens were modelled. In particular, relationships between the occurrence of alien plants and (1) elevation or (2) the climatic variables, were tested by applying generalised linear models and generalised linear mixed models; the model parameters obtained were used to estimate upper elevation limits of alien occurrence and their related climate values. Sixty-eight alien species were encountered, the majority (71%) invasive in Italy and worldwide. A steep decrease in alien species richness with elevation was found, with the probability of alien species occurrence decreasing by half for each 100 m increase in elevation. Minimal adequate models based on (1) non-transformed climatic variables and (2) derived PCA values, confirmed that occurrence of alien plant species along the elevation gradient was positively related to the minimum temperature, the mean temperature and the heat sum for the spring season, rather than to the incidence of absolute minimum temperature and frost days, as usually assumed. Although further experimental analyses are needed, these results support the hypothesis that, referring to climate factors, elevation limits along rivers are mainly established by low spring temperatures which operate at the level of population viability rather than plant survival.     

Co-invasive exotic pines and their ectomycorrhizal symbionts show capabilities for wide distance and altitudinal range expansion

We asked if exotic Pinus elliotti seedlings can survive and form ectomycorrhizas at higher elevations and long distances from their current range, and which ECM partners disperse to these soils. We selected three plots at four grassland sites along an altitudinal gradient (900, 1600, 2200, and 2700 m asl) established at c. 110, 3000, 6000, and 9000 m from the closest pine plantation, respectively. We combined field experiments with glasshouse assays to assess survival and ECM fungi in roots and soils. A pine plantation close to the lowest site was also selected for DNA metabarcoding of soils. Pine seedlings survived at all altitudes but not all formed mycorrhizas. They formed mycorrhizas with Suillus granulatus at 900, 1600, and 2200 m asl (i.e. up to 6000 m from the closest pine plantation), and with Rhizopogon pseudoroseolus and Thelephora terrestris at lower altitudes and distances. Twelve ECM fungal OTUs were found in grasslands and 34 were detected in the pine plantation. Although richness and abundance of ECM fungi decreased with increasing distance from the pine plantation, there was at least one non-native ECM fungal species present in each sampling site, even at 2700 masl and 9000 m distance from the closest plantation. This study provides evidence that the availability of suitable fungal symbionts might constrain but not hinder the expansion of a pine species over wide distances and altitudinal zones even in areas with no native ECM fungi.     

Nest‐site selection by Interior Least Terns and Piping Plovers at managed,off‐channel sites along the Central Platte River in Nebraska,USA

Given the high productivity of Interior Least Terns (Sternula antillarum athalassos) and Piping Plovers (Charadrius melodus) on constructed off‐channel nesting sites along the central Platte River in Nebraska, USA, and the possibility of creating similar habitats at other locations in their breeding range, understanding how these species use off‐channel nesting habitats is important. We used data collected along the central Platte River in Nebraska, USA, over a 15‐year period (2001–2015), and a discrete‐choice modeling framework to assess the effects of physical site attributes and inter‐ and intraspecific associations on off‐channel nest‐site selection by Interior Least Terns and Piping Plovers. We found that Piping Plovers avoided nesting near each other, whereas colonial Interior Least Terns selected nest sites near those of conspecifics. In addition, the relative probability of use for both species was maximized when distance to the nearest predator perch was ≥ 150 m and elevation above the waterline was ≥ 3 m. Probability of use for nesting by Interior Least Terns increased as distance to water increased, whereas the probability of use by Piping Plovers was maximized when distance to water was ~50 m. Our results suggest that important features of constructed, off‐channel nesting sites for both species should include no potential predator perches within 150 m of nesting habitat and nesting areas at least 3 m above the waterline. Efficient site designs for Interior Least Terns would be circular, maximizing the area of nesting habitat away from the shoreline, whereas an effective site design for Piping Plovers would be more linear, maximizing the area of nesting habitat near the waterline. An efficient site design for both species would be lobate, incorporating centralized nesting habitat for Interior Least Terns and increased access to foraging areas for nesting and brood‐rearing Piping Plovers.     

Living at the edge: lower success of eggs and hatchlings at lower elevation may shape range limits in an alpine lizard

Studies on range limits clarify the factors involved in the extent of species occurrence and shed light on the limits to adaptation. We studied the effects of elevational variation on the thermal dependence of fitness‐related traits (incubation time, hatching rate, and survivorship, size, and condition of hatchlings) to assess the role of incubation requirements in distribution range limits of the alpine endemic Iberolacerta cyreni. We captured gravid females from two core (summit) and two marginal (low‐elevation edge) populations, we incubated their eggs at three temperatures (22, 26, and 30 °C), and we monitored phenotypic effects. Viability of eggs and hatchlings decreased, independently of elevation, as incubation temperature increased. Hatching success and embryo survivorship were lower for clutches from low‐elevation areas than for those from mountain summits, showing that lizards face difficulties thriving at the low‐elevation edge of their range. Such difficulties were partly counterbalanced by faster postnatal growth at lower elevations, leading to increased adult size and higher fecundity. High incubation temperature had detrimental effects also at low‐elevation areas, and no elevational variation in the thermal dependence of hatchling traits was detected. We suggest that temperature effects on egg development and the lack of selective pressures strong enough to foster local adaptation at marginal areas, combined with extended egg retention, may contribute to shape the range limits of these alpine oviparous reptiles.