Predictable Variation of Range-Sizes across an Extreme Environmental Gradient in a Lizard Adaptive Radiation: Evolutionary and Ecological Inferences

Large-scale patterns of current species geographic range-size variation reflect historical dynamics of dispersal and provide insights into future consequences under changing environments. Evidence suggests that climate warming exerts major damage on high latitude and elevation organisms, where changes are more severe and available space to disperse tracking historical niches is more limited. Species with longer generations (slower adaptive responses), such as vertebrates, and with restricted distributions (lower genetic diversity, higher inbreeding) in these environments are expected to be particularly threatened by warming crises. However, a well-known macroecological generalization (Rapoport's rule) predicts that species range-sizes increase with increasing latitude-elevation, thus counterbalancing the impact of climate change. Here, I investigate geographic range-size variation across an extreme environmental gradient and as a function of body size, in the prominent Liolaemus lizard adaptive radiation. Conventional and phylogenetic analyses revealed that latitudinal (but not elevational) ranges significantly decrease with increasing latitude-elevation, while body size was unrelated to range-size. Evolutionarily, these results are insightful as they suggest a link between spatial environmental gradients and range-size evolution. However, ecologically, these results suggest that Liolaemus might be increasingly threatened if, as predicted by theory, ranges retract and contract continuously under persisting climate warming, potentially increasing extinction risks at high latitudes and elevations

Intraspecific predation in the Liolaemus lizard radiation: a primer

Intraspecific predation (cannibalism) occurs in a number of lizard species, although the incidence of this trophic behaviour appears to be infrequent. Within the South American Liolaemus lizard evolutionary radiation, the primary component of the carnivorous diet (herbivory has also evolved in several species) includes a diversity of arthropods and other invertebrates. However, a number of cases of saurophagy, mostly cannibalism, have also been reported in different species. Here, I review for the first time the occurrence of cannibalism in this prominent group of lizards, one of the largest and more ecologically diverse living adaptive radiations. Adults of five Liolaemus species have previously been reported to feed on conspecific hatchlings, and one additional case (L. zapallarensis) is reported in this paper. In general, cannibalism is found in species located at the right side of the body size frequency distribution, while no cases have been reported on the far left range of body sizes, suggesting that cannibalism is more common in large species. Only one species of medium size (L. darwinii) is known to feed on conspecifics. In addition, males are responsible for 83% of the cases of cannibalism, while only one case of female cannibalism is known (L. chiliensis). Finally, no clear phylogenetic signal can be inferred from these observations, as the six reported cannibal species belong to four of the five main Liolaemus clades

Heterogeneous tempo and mode of evolutionary diversification of compounds in lizard chemical signals

Important part of the multivariate selection shaping social and interspecific interactions among and within animal species emerges from communication. Therefore, understanding the diversification of signals for animal communication is a central endeavor in evolutionary biology. Over the last decade, the rapid development of phylogenetic approaches has promoted a stream of studies investigating evolution of communication signals. However, comparative research has primarily focused on visual and acoustic signals, while the evolution of chemical signals remains largely unstudied. An increasing interest in understanding the evolution of chemical communication has been inspired by the realization that chemical signals underlie some of the major interaction channels in a wide range of organisms. In lizards, in particular, chemosignals play paramount roles in female choice and male–male competition, and during community assembly and speciation. Here, using phylogenetic macro-evolutionary modeling, we show for the very first time that multiple compounds of scents for communication in lizards have diversified following highly different evolutionary speeds and trajectories. Our results suggest that cholesterol, α-tocopherol, and cholesta-5,7-dien-3-ol have been subject to stabilizing selection (Ornstein–Uhlenbeck model), whereas the remaining compounds are better described by Brownian motion modes of evolution. Additionally, the diversification of the individual compounds has accumulated substantial relative disparity over time. Thus, our study reveals that the chemical components of lizard chemosignals have proliferated across different species following compound-specific directions

The macroecology of chemical communication in lizards: do climatic factors drive the evolution of signalling glands?

Chemical communication plays a pivotal role in shaping sexual and ecological interactions among animals. In lizards, fundamental mechanisms of sexual selection such as female mate choice have rarely been shown to be influenced by quantitative phenotypic traits (e.g., ornaments), while chemical signals have been found to potentially influence multiple forms of sexual and social interactions, including mate choice and territoriality. Chemical signals in lizards are secreted by glands primarily located on the edge of the cloacae (precloacal glands, PG) and thighs (femoral glands), and whose interspecific and interclade number ranges from 0 to >100. However, elucidating the factors underlying the evolution of such remarkable variation remains an elusive endeavour. Competing hypotheses suggest a dominant role for phylogenetic conservatism (i.e., species within clades share similar numbers of glands) or for natural selection (i.e., their adaptive diversification results in deviating numbers of glands from ancestors). Using the prolific Liolaemus lizard radiation from South America (where precloacal glands vary from 0-14), we present one of the largest-scale tests of both hypotheses to date. Based on climatic and phylogenetic modelling, we show a clear role for both phylogenetic inertia and adaptation underlying gland variation: (i) solar radiation, net primary productivity, topographic heterogeneity and precipitation range have a significant effect on number of PG variation, (ii) humid and cold environments tend to concentrate species with a higher number of glands, (iii) there is a strong phylogenetic signal that tends to conserve the number of PG within clades. Collectively, our study confirms that the inertia of niche conservatism can be broken down by the need of species facing different selection regimes to adjust their glands to suit the demands of their specific environments

A quantitative analysis of objective feather color assessment: Measurements in the laboratory do not reflect true plumage color

© 2016 American Ornithologists' Union.An important driver of the evolution of animal coloration is sexual selection operating on traits that are used to transmit information to rivals and potential mates, which has a major impact on fitness. Reflectance spectrometry has become a standard color-measuring tool, especially after the discovery of tetrachromacy in birds and their ability to detect UV light. Birds' plumage patterns may be invisible to humans, and therefore the establishment of reliable and quantitatively objective ways of assessing coloration not dependent on human vision is a technical need of primary importance. Plumage coloration measurements can be taken directly on live birds in the field, or in the laboratory (e.g., on collected feathers). However, which of these 2 approaches offers a more reliable, repeatable sampling method remains an unsolved question. Using a spectrophotometer, we measured melanin-based coloration in the plumage of Barn Swallows (Hirundo rustica). We assessed the repeatability of measures obtained with both traditional sampling methods to quantitatively determine their reliability. We used an ANOVA-based method for calculating the repeatability of measurements from 2 years separately, and a GLMM-based method to calculate overall adjusted repeatabilities for both years. The results of our study indicate a great disparity between color measurements obtained using both sampling methods and a low comparability across them. Assuming that measurements taken in the field reflect the real or "true" color of plumage, we may conclude that there is a lack of reliability of the laboratory method to reflect this true color in melanin-based plumages. Likewise, we recommend the use of the GLMM-based statistical method for repeatability calculations, as it allows the inclusion of random factors and the calculation of more realistic, adjusted repeatabilities. It also reduces the number of necessary tests, thereby increasing power, and it allows easy calculation of 95% CIs, a measure of the reliability and precision of effect-size calculations.published_or_final_versio

The multiple origins of sexual size dimorphism in global amphibians

Aim: Body size explains most variation in fitness within animal populations and is therefore under constant selection from ecological and reproductive pressures which often promote its evolution in sex-specific directions, leading to sexual size dimorphism (SSD). Multiple hypotheses have been proposed to explain the vast diversity of SSD across species. These hypotheses emphasise (i) the mate competition benefits to larger male size (sexual selection), (ii) the benefits of larger female size for fecundity (fecundity selection), (iii) the simultaneous benefits of niche divergence for males and females to reduce inter-sexual competition for ecological resources (natural selection), and (iv) the underlying impact of geographic variation in climatic pressures expected to shape large-scale patterns of SSD in synergy with the above selection pressures (e.g., intensification of fecundity selection as breeding seasons shorten). Based on a novel, global-scale amphibian dataset, we address the shortage of large-scale, integrative tests of these four hypotheses. Location: Global. Time period: Extant. Major taxa studied: Class Amphibia. Methods: Using a >3,500 species dataset spanning body size, ecological, life history, geographic and climatic data, we performed phylogenetic linear models to address the sexual, fecundity, ecological and climatic hypotheses of SSD. Results: SSD evolution is discordant between anurans and salamanders. Anuran SSD is shaped by climate – male-biased SSD increases with temperature seasonality – and by nesting site. In salamanders, SSD converges across species that occupy the same microhabitat types (‘ecodimorphs’), while reproductive or climatic pressures have no effects on their SSD. These contrasts are associated with latitudinal gradients of SSD in anurans, but not in salamanders. Main conclusions: Amphibian SSD is driven by ecological and climatic pressures, while no roles for sexual or fecundity selection were detected. We show that macroevolutionary processes determined by different forms of selection lead to latitudinal patterns of trait diversity, and the lack of them

Global Protected Areas as refuges for amphibians and reptiles under climate change

Protected Areas (PAs) are the cornerstone of biodiversity conservation. Here, we collated distributional data for >14,000 (~70% of) species of amphibians and reptiles (herpetofauna) to perform a global assessment of the conservation effectiveness of PAs using species distribution models. Our analyses reveal that >91% of herpetofauna species are currently distributed in PAs, and that this proportion will remain unaltered under future climate change. Indeed, loss of species’ distributional ranges will be lower inside PAs than outside them. Therefore, the proportion of effectively protected species is predicted to increase. However, over 7.8% of species currently occur outside PAs, and large spatial conservation gaps remain, mainly across tropical and subtropical moist broadleaf forests, and across non-high-income countries. We also predict that more than 300 amphibian and 500 reptile species may go extinct under climate change over the course of the ongoing century. Our study highlights the importance of PAs in providing herpetofauna with refuge from climate change, and suggests ways to optimize PAs to better conserve biodiversity worldwide

Global protected areas as refuges for amphibians and reptiles under climate change

Protected Areas (PAs) are the cornerstone of biodiversity conservation. Here, we collated distributional data for >14,000 (~70% of) species of amphibians and reptiles (herpetofauna) to perform a global assessment of the conserva- tion effectiveness of PAs using species distribution models. Our analyses reveal that >91% of herpetofauna species are currently distributed in PAs, and that this proportion will remain unaltered under future climate change. Indeed, loss of species’ distributional ranges will be lower inside PAs than outside them. Therefore, the proportion of effectively protected species is predicted to increase. However, over 7.8% of species currently occur outside PAs, and large spatial conservation gaps remain, mainly across tropical and subtropical moist broadleaf forests, and across non-high-income countries. We also predict that more than 300 amphibian and 500 reptile species may go extinct under climate change over the course of the ongoing century. Our study highlights the importance of PAs in providing herpetofauna with refuge from climate change, and suggests ways to optimize PAs to better conserve biodiversity worldwide

Field observations on the natural history and breeding behavior of the Atacama toad (Rhinella atacamensis, Anura: Bufonidae) from Chile

Amphibians have diversified predominantly across tropical environments where humidity, temperature and microhabitat availability facilitate demographic stability. However, a number of lineages have colonized extreme deserts, where their diversities are considerably lower. One species in particular, the Atacama toad (Rhinella atacamensis), has adapted to the Atacama Desert, the driest place on Earth. Despite the ecological uniqueness of this species, most aspects of its natural history, reproduction, patterns of activity and behavior remain unknown. Using camera traps and in situ field observations, we report a set of novel natural history findings in a population from the Llanos de Challe National Park. We show that R. atacamensis remains hidden from the sun in small pools under vegetation, where 100% of observed femalJes have a male permanently attached in amplexus at all times. The toads emerge to gather in small ponds after the sunset (~20:00h), where males engage in active contests over females, with up to four males competing for one female. They retreat into the covered pools after ~06:00h. Eggs laid during the night hatch within 24 hours. We discuss these observations in the context of the global diversity of bufonid toads in general, and in relation with the effects that selection emerging from the harsh conditions of Atacama may exert on the expression of these traits

Ant assemblages have darker and larger members in cold environments

Aim In ectotherms, the colour of an individual's cuticle may have important thermoregulatory and protective consequences. In cool environments, ectotherms should be darker, to maximize heat gain, and larger, to minimize heat loss. Dark colours should also predominate under high UV‐B conditions because melanin offers protection. We test these predictions in ants (Hymenoptera: Formicidae) across space and through time based on a new, spatially and temporally explicit, global‐scale combination of assemblage‐level and environmental data. Location Africa, Australia and South America. Methods We sampled ant assemblages (n = 274) along 14 elevational transects on three continents. Individual assemblages ranged from 250 to 3000 m a.s.l. (minimum to maximum range in summer temperature of 0.5–35 °C). We used mixed‐effects models to explain variation in assemblage cuticle lightness. Explanatory variables were average assemblage body size, temperature and UV‐B irradiation. Annual temporal changes in lightness were examined for a subset of the data. Results Assemblages with large average body sizes were darker in colour than those with small body sizes. Assemblages became lighter in colour with increasing temperature, but darkened again at the highest temperatures when there were high levels of UV‐B. Through time, temperature and body size explained variation in lightness. Both the spatial and temporal models explained c. 50% of the variation in lightness. Main conclusions Our results are consistent with the thermal melanism hypothesis, and demonstrate the importance of considering body size and UV‐B radiation exposure in explaining the colour of insect cuticle. Crucially, this finding is at the assemblage level. Consequently, the relative abundances and identities of ant species that are present in an assemblage can change in accordance with environmental conditions over elevation, latitude and relatively short time spans. These findings suggest that there are important constraints on how ectotherm assemblages may be able to respond to rapidly changing environmental conditions