Search

Synopsis Efficient comparisons of biological color patterns are critical for understanding the mechanisms by which organisms evolve in nature, including sexual selection, predator–prey interactions, and thermoregulation. However, limbless, elongate, or spiral-shaped organisms do not conform to the standard orientation and photographic techniques required for many automated analyses. Currently, large-scale color analysis of elongate animals requires time-consuming manual landmarking, which reduces their representation in coloration research despite their ecological importance. We present Batch-Mask: an automated, customizable workflow to automatically analyze large photographic datasets to isolate non-standard biological organisms from the background. Batch-Mask is completely open-source and does not depend on any proprietary software. We also present a user guide for fine-tuning weights to a custom dataset and incorporating existing manual visual analysis tools (e.g., micaToolbox) into a single automated workflow for comparing color patterns across images. Batch-Mask was 60x faster than manual landmarking and produced masks that correctly identified 96% of all snake pixels. To validate our approach, we used micaToolbox to compare pattern energy in a sample set of snake photographs segmented by Batch-Mask and humans and found no significant difference in the output results. The fine-tuned weights, user guide, and automated workflow substantially decrease the amount of time and attention required to quantitatively analyze non-standard biological subjects. With these tools, biologists can compare color, pattern, and shape differences in large datasets that include significant morphological variation in elongate body forms. This advance is especially valuable for comparative analyses of natural history collections across a broad range of morphologies. Through landmark-free automation, Batch-Mask can greatly expand the scale of space, time, or taxonomic breadth across which color variation can be quantitatively examined.

Diffusible iodine‐based contrast‐enhanced computed tomography (diceCT) visualizes soft tissue from micro‐CT (µCT) scans of specimens to uncover internal features and natural history information without incurring physical damage via dissection. Unlike hard‐tissue imaging, taxonomic sampling within diceCT datasets is currently limited. To initiate best practices for diceCT in a nonmodel group, we outline a guide for staining and high‐throughput µCT scanning in snakes. We scanned the entire body and one region of interest (i.e., head) for 23 specimens representing 23 species from the clades Aniliidae, Dipsadinae, Colubrinae, Elapidae, Lamprophiidae, and Viperidae. We generated 82 scans that include 1.25% Lugol's iodine stained (soft tissue) and unstained (skeletal) data for each specimen. We found that duration of optimal staining time increased linearly with body size; head radius was the best indicator. Postreconstruction of scans, optimal staining was evident by evenly distributed grayscale values and clear differentiation among soft‐tissue anatomy. Under and over stained specimens produced poor contrast among soft tissues, which was often exacerbated by user bias during “digital dissections” (i.e., segmentation). Regardless, all scans produced usable data from which we assessed a range of downstream analytical applications within ecology and evolution (e.g., predator‐prey interactions, life history, and morphological evolution). Ethanol destaining reversed the known effects of iodine on the exterior appearance of physical specimens, but required substantially more time than reported for other destaining methods. We discuss the feasibility of implementing diceCT techniques for a new user, including approximate financial and temporal commitments, required facilities, and potential effects of staining on specimens. We present the first high‐throughput workflow for full‐body skeletal and diceCT scanning in snakes, which can be generalized to any elongate vertebrates, and increases publicly available diceCT scans for reptiles by an order of magnitude., We present the first high‐throughput workflow for full‐body skeletal and soft‐tissue (dice) CT scans in preserved snakes. These methods increase publicly available diceCT scans for reptiles by an order of magnitude and can be generalized to any elongate vertebrates.

Animals in nature use many strategies to evade or deter their predators. Within venomous snake mimicry, stereotyped anti-predator behaviours are hypothesized to be effective warning signals under strong selection for independent convergence across species. However, no studies have systematically quantified snake anti-predator displays across taxonomically broad clades to examine how these behaviours evolve within a comparative methods framework. Here we describe a new high-throughput approach for collecting and quantifying anti-predator displays in snakes that demonstrates both low observer bias and infinite extension. Then, we show this method’s utility by comparing 20 species spanning six taxonomic families from Peru. We found that a simple experimental set-up varying simulated predator cues was successful in eliciting displays across species and that high-speed videography captured a great diversity of anti-predator responses. Although display components show complicated patterns of covariance, we found support for behavioural convergence in anti-predator displays among elapid coral snakes and their distantly related mimics. Our approach provides new analytical opportunities for both behaviour and kinematics, especially macroevolutionary analyses across clades with similar difficulty in observing or comparing trait diversity.

Efficient comparisons of biological color patterns are critical for understanding the mechanisms by which organisms evolve in ecosystems, including sexual selection, predator-prey interactions, and thermoregulation. However, elongate or spiral-shaped organisms do not conform to the standard orientation and photographic techniques required for automated analysis. Currently, large-scale color analysis of elongate animals requires time-consuming manual landmarking, which reduces their representation in coloration research despite their ecological importance.We present Batch-Mask: an automated and customizable workflow to facilitate the analysis of large photographic data sets of non-standard biological subjects. First, we present a user guide to run an open-source region-based convolutional neural network with fine-tuned weights for identifying and isolating a biological subject from a background (masking). Then, we demonstrate how to combine masking with existing manual visual analysis tools into a single streamlined, automated workflow for comparing color patterns across images.Batch-Mask was 60x faster than manual landmarking, produced masks that correctly identified 96% of all snake pixels, and produced pattern energy results that were not significantly different from the manually landmarked data set.The fine-tuned weights for the masking neural network, user guide, and automated workflow substantially decrease the amount of time and attention required to quantitatively analyze non-standard biological subjects. By using these tools, biologists will be able to compare color, pattern, and shape differences in large data sets that include significant morphological variation in elongate body forms. This advance will be especially valuable for comparative analyses of natural history collections, and through automation can greatly expand the scale of space, time, or taxonomic breadth across which color variation can be quantitatively examined.

Gut microbiomes perform essential services for their hosts, including helping them to digest food and manage pathogens and parasites. Performing these services requires a diverse and constantly changing set of metabolic functions from the bacteria in the microbiome. The metabolic repertoire of the microbiome is ultimately dependent on the outcomes of the ecological interactions of its member microbes, as these interactions in part determine the taxonomic composition of the microbiome. The ecological processes that underpin the microbiome's ability to handle a variety of metabolic challenges might involve rapid turnover of the gut microbiome in response to new metabolic challenges, or it might entail maintaining sufficient diversity in the microbiome that any new metabolic demands can be met from an existing set of bacteria. To differentiate between these scenarios, we examine the gut bacteria and resident eukaryotes of two generalist‐insectivore lizards, while simultaneously identifying the arthropod prey each lizard was digesting at the time of sampling. We find that the cohorts of bacteria that occur significantly more or less often than expected with arthropod diet items or eukaryotes include bacterial species that are highly similar to each other metabolically. This pattern in the bacterial microbiome could represent an early step in the taxonomic shifts in bacterial microbiome that occur when host lineages change their diet niche over evolutionary timescales., The ecological processes that underpin the gut microbiome's metabolic response might involve rapid turnover of the gut microbiome in response to new metabolic challenges, or it might entail maintaining sufficient diversity in the microbiome that any new metabolic demands can be met from an existing set of bacteria. We find that the cohorts of bacteria that occur significantly more or less often than expected with arthropod diet items or eukaryotes in lizard hosts include fewer bacterial species than would be expected by chance, but that those species are highly similar to each other metabolically.

Covariation among traits shapes both phenotypic evolution and ecological interactions across space and time. However, rampant geographical variation in the strength and direction of such correlations can be particularly difficult to explain through generalized mechanisms. By integrating population genomics, surveys of natural history collections and spatially explicit analyses, we tested multiple drivers of trait correlations in a coral snake mimic that exhibits remarkable polymorphism in mimetic and non-mimetic colour traits. We found that although such traits co-occur extensively across space, correlations were best explained by a mixture of genetic architecture and correlational selection, rather than by any single mechanism. Our findings suggest that spatially complex trait distributions may be driven more by the simple interaction between multiple processes than by complex variation in one mechanism alone. These interactions are particularly important in mimicry systems, which frequently generate striking geographical variation and genetic correlations among colour pattern traits.

This paper presents a soft-robotic platform for exploring the ecological relevance of non-locomotory movements via animal-robot interactions. Coral snakes (genus Micrurus) and their mimics use vigorous, non-locomotory, and arrhythmic thrashing to deter predation. There is variation across snake species in the duration and curvature of anti-predator thrashes, and it is unclear how these aspects of motion interact to contribute to snake survival. In this work, soft robots composed of fiber-reinforced elastomeric enclosures (FREEs) are developed to emulate the anti-predator behaviors of three genera of snake. Curvature and duration of motion are estimated for both live snakes and robots, providing a quantitative assessment of the robots' ability to emulate snake poses. The curvature values of the fabricated soft-robotic head, midsection, and tail segments are found to overlap with those exhibited by live snakes. Soft robot motion durations were less than or equal to those of snakes for all three genera. Additionally, combinations of segments were selected to emulate three specific snake genera with distinct anti-predatory behavior, producing curvature values that aligned well with live snake observations., 6 pages, 7 figures

Background Fangs are a putative key innovation that revolutionized prey capture and feeding in snakes, and – along with their associated venom phenotypes – have made snakes perhaps the most medically-significant vertebrate animals. Three snake clades are known for their forward-positioned fangs, and these clades (Elapidae, Viperidae, and Atractaspidinae) contain the majority of snakes that are traditionally considered venomous. However, many other snakes are “rear-fanged”: they possess potentially venom-delivering teeth situated at the rear end of the upper jaw. Quantification of fang phenotypes – and especially those of rear-fanged species – has proved challenging or impossible owing to the small size and relative rarity of many such snakes. Consequently, it has been difficult to understand the evolutionary history of both venom and prey-capture strategies across extant snakes. We quantified variation in the dentition of 145 colubriform (“advanced”) snake species using microCT scanning and compared dental characters with ecological data on species’ diet and prey capture method(s) to understand broader patterns in snake fang evolution. Results Dental traits such as maxilla length, tooth number, and fang size show strong phylogenetic signal across Colubriformes. We find extreme heterogeneity and evolutionary lability in the rear-fanged phenotype in colubrid (colubrine, dipsadine, and natricine lineages) and lamprophiid snakes, in contrast to relative uniformity in the front fanged phenotypes of other groups (vipers and, to a lesser extent, elapids). Fang size and position are correlated with venom-use in vipers, elapids, and colubrid snakes, with the latter group shifting fangs anteriorly by shortening the entire maxillary bone. We find that maxilla length and tooth number may also be correlated with the evolution of dietary specialization. Finally, an ancestral state reconstruction suggests that fang loss is a recurring phenomenon in colubrid snakes, likely accompanied by shifts in diet and prey capture mode. Conclusions Our study provides a framework for quantifying the complex morphologies associated with venom use in snakes. Our results suggest that fang phenotypes, and particularly the rear-fanged phenotype, in snakes are both diverse and labile, facilitating a wide range of ecological strategies and contributing to spectacular radiations of these organisms in tropical and subtropical biomes worldwide.

Delimiting species is a crucial goal of integrative biology, and yet can be misled by homoplasy and high levels of morphological variation. The snake tribe Sonorini contains three genera that have long confounded taxonomists: Chilomeniscus, Chionactis and Sonora. Dynamic colour evolution in this group, including rampant geographic variation in colour and colour polymorphism, has led to a chaotic taxonomy. We used mitochondrial and high-throughput nuclear data (ddRADseq) and complete taxonomic sampling of each genus to reconstruct phylogenetic relationships and systematically revise the genus. Our research revealed that Sonora is paraphyletic with regards to Chilomeniscus and Chionactis and that at least one species (S. semiannulata) is paraphyletic with respect to at least one other recognized species. Additionally, we found substantial undescribed genetic diversity within multiple species which is incongruent with morphological variation in coloration. Accordingly, we proposed synonymizing Chiona...

Color polymorphism in natural populations can manifest as a striking patchwork of phenotypes in space, with neighboring populations characterized by dramatic differences in morph composition. These geographic mosaics can be challenging to explain in the absence of localized selection because they are unlikely to result from simple isolation-by-distance or clinal variation in selective regimes. To identify processes that can lead to the formation of geographic mosaics, we developed a simulation-based model to explore the influence of predator perspective, selection, migration, and genetic linkage of color loci on allele frequencies in polymorphic populations over space and time. Using simulated populations inspired by the biology of Heliconius longwing butterflies, Cepaea land snails, Oophaga poison frogs, and Sonora ground snakes, we found that the relative sizes of predator and prey home ranges can produce large differences in morph composition between neighboring populations under both positive and negative frequency-dependent selection. We also demonstrated the importance of the interaction of predator perspective with the type of frequency dependence and localized directional selection across migration and selection intensities. Our results show that regional-scale predation can promote the formation of phenotypic mosaics in prey species, without the need to invoke spatial variation in selective regimes. We suggest that predator behavior can play an important and underappreciated role in the formation and maintenance of geographic mosaics in polymorphic species.

Our knowledge of ecological interactions that bolster ecosystem function and productivity has broad applications to the management of agricultural systems. Studies suggest that the presence of generalist predators in agricultural landscapes leads to a decrease in the abundance of herbivorous pests, but our understanding of how these interactions vary across taxa and along gradients of management intensity and eco‐geographic space remains incomplete. In this study, we assessed the functional response and biocontrol potential of a highly ubiquitous insectivore (lizards in the genus Anolis) on the world's most important coffee pest, the coffee berry borer (Hypothalemus hampei). We conducted field surveys and laboratory experiments to examine the impact of land‐use intensification on species richness and abundance of anoles and the capacity of anoles to reduce berry borer infestations in mainland and island coffee systems. Our results show that anoles significantly reduce coffee infestation rates in laboratory settings (Mexico, p = .03, F = 5.13 df = 1, 35; Puerto Rico, p = .014, F = 8.82, df = 1, 10) and are capable of consuming coffee berry borers in high abundance. Additionally, diversified agroecosystems bolster anole abundance, while high‐intensity practices, including the reduction of vegetation complexity and the application of agrochemicals were associated with reduced anole abundance. The results of this study provide supporting evidence of the positive impact of generalist predators on the control of crop pests in agricultural landscapes, and the role of diversified agroecosystems in sustaining both functionally diverse communities and crop production in tropical agroecosystems.

The lowland rainforests of the Amazon basin harbor some of the most species-rich reptile communities on Earth. However, there is considerable heterogeneity among climatically-similar sites across the Amazon basin, and faunal surveys for southwestern Amazonia in particular have revealed lower species diversity relative to sites in the northwestern and central Amazon. Here, we report a herpetofaunal inventory for Los Amigos Biological Station (LABS), a lowland site located in the Madre de Dios watershed of southern Peru. By combining active search and passive trapping methods with prior records for the site, we provide a comprehensive species list for squamate reptiles from LABS. We also estimate an &ldquo, expected&rdquo, list for LABS by tabulating additional taxa known from the regional species pool that we consider to have a high probability of detection with further sampling. The LABS total of 60 snake and 26 lizard taxa is perhaps the highest for any single site in the southern Amazon. Our estimate of the regional species pool for LABS suggests that the southwestern Amazonian lowlands harbor at least 25% fewer species of snakes relative to the western equatorial Amazon, a diversity reduction that is consistent with patterns observed in several other taxonomic groups. We discuss potential causes of this western Amazonian richness gradient and comment on the relationship between spatial diversity patterns in squamates and other taxa in the Amazon basin.

Warning signals in chemically defended organisms are critical components of predator-prey interactions, often requiring multiple coordinated display components for a signal to be effective. When threatened by a predator, venomous coral snakes (genusMicrurus) display a vigorous, non-locomotory thrashing behaviour that has been only qualitatively described. Given the high-contrast and often colourful banding patterns of these snakes, this thrashing display is hypothesized to be a key component of a complex aposematic signal under strong stabilizing selection across species in a mimicry system.By experimentally testing snake response across simulated predator cues, we analysed variation in the presence and expression of a thrashing display across five species of South American coral snakes.Although the major features of the thrash display were conserved across species, we found significant variation in the propensity to perform a display at all, the duration of thrashing, and the curvature of snake bodies that was mediated by predator cue type, snake body size, and species identity. We also found an interaction between curve magnitude and body location that clearly shows which parts of the display vary most across individuals and species.Our results suggest that contrary to the assumption in the literature that all species and individuals perform the same display, a high degree of variation persists in thrashing behaviour exhibited byMicruruscoral snakes despite presumably strong selection to converge on a common signal. This quantitative behavioural characterization presents a new framework for analysing the non-locomotory motions displayed by snakes in a broader ecological context, especially for signalling systems with complex interaction across multiple modalities.

Procedure for multilocus sequence typing (MLST) from amphibian skin swabs using Sanger sequencing.

Protocol for extracting DNA from amphibian skin swabs using the Qiagen DNeasy blood and Tissue Kit.

Method for quantitative (real-time) PCR using TaqMan reagents

Protocol for extracting DNA from amphibian skin swabs using the Qiagen DNeasy blood and Tissue Kit.

Genetic structure can be influenced by local adaptation to environmental heterogeneity and biogeographic barriers, resulting in discrete population clusters. Geographic distance among populations, however, can result in continuous clines of genetic divergence that appear as structured populations. Here, we evaluate the relevant importance of these three factors over a landscape characterized by environmental heterogeneity and the presence of a hypothesized biogeographic barrier in producing population genetic structure within 13 codistributed snake species using a genomic data set. We demonstrate that geographic distance and environmental heterogeneity across western North America contribute to population genomic divergence. Surprisingly, landscape features long thought to contribute to biogeographic barriers play little role in divergence community wide. Our results suggest that isolation by environment is the most important contributor to genomic divergence. Furthermore, we show that models of population clustering that incorporate spatial information consistently outperform nonspatial models, demonstrating the importance of considering geographic distances in population clustering. We argue that environmental and geographic distances as drivers of community-wide divergence should be explored before assuming the role of biogeographic barriers.

Genetic polymorphism in males has long been considered paradoxical because sexual selection is expected to deplete additive genetic variation. Although studies have shown how divergent selection between populations can make that work, it is very rare to find disruptive selection within one population. Since intersexual selection can have a significant effect on the phenotypic morphospace of the opposite sex, we analysed the role of female preference as a disruptive selective force. In this study we evaluated how female preference acts on anatomical and sex-related behavioural traits of two male morphs in the jumping spider Maevia inclemens. We used mate choice trials to analyse how female preference for the two morphs varied. The tests indicated that females preferred opposite values of two anatomical and two behavioural traits for each male morph. This study is, to our knowledge, the first to show disruptive sexual selection in Arachnidae and significantly expands the realm of disruptive selection, by adding one more case to the very few documented instances. These processes, which act entirely within a species, are of particular interest because they could contribute to the evolution of reproductive isolation and sympatric speciation by sexual selection, a controversial topic in evolutionary biology.

Background Reduced representation genomic datasets are increasingly becoming available from a variety of organisms. These datasets do not target specific genes, and so may contain sequences from parasites and other organisms present in the target tissue sample. In this paper, we demonstrate that (1) RADseq datasets can be used for exploratory analysis of tissue-specific metagenomes, and (2) tissue collections house complete metagenomic communities, which can be investigated and quantified by a variety of techniques. Methods We present an exploratory method for mining metagenomic “bycatch” sequences from a range of host tissue types. We use a combination of the pyRAD assembly pipeline, NCBI’s blastn software, and custom R scripts to isolate metagenomic sequences from RADseq type datasets. Results When we focus on sequences that align with existing references in NCBI’s GenBank, we find that between three and five percent of identifiable double-digest restriction site associated DNA (ddRAD) sequences from host tissue samples are from phyla to contain known blood parasites. In addition to tissue samples, we examine ddRAD sequences from metagenomic DNA extracted snake and lizard hind-gut samples. We find that the sequences recovered from these samples match with expected bacterial and eukaryotic gut microbiome phyla. Discussion Our results suggest that (1) museum tissue banks originally collected for host DNA archiving are also preserving valuable parasite and microbiome communities, (2) that publicly available RADseq datasets may include metagenomic sequences that could be explored, and (3) that restriction site approaches are a useful exploratory technique to identify microbiome lineages that could be missed by primer-based approaches.

Outbreaks of emerging infectious diseases are becoming more frequent as climate changes wildlife communities at unprecedented rates, driving population declines and raising concerns for species conservation. One critical disease is the global pandemic of chytridiomycosis in frogs, which can be caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd). Although there is clear evidence for Bd-induced mortality across high-elevation frog communities, little attention is given to the role of lowlands in Bd’s persistence and spread because low elevations are assumed to be too warm to harbor significant levels of Bd. Here, we report widespread Bd infection across 80 frog species from three sites in the lowland Peruvian Amazon, an area with no documented Bd-related amphibian declines. Despite observing no clinical signs of infection in the field, we found that 24–46% of individuals were infected per site (up to ≈105,000 zoospore equivalents per frog) by three Bd strains from the global pandemic lineage (Bd-GPL). We also found collection site and seasonal effects to be only weak predictors of Bd prevalence and load, with lower elevation and drier habitats marginally decreasing both prevalence and load. We found no further effect of host phylogeny, ecotype, or body size. Our results showing high and widespread prevalence across a lowland tropical ecosystem contradict the expectations based on the global pattern of pathogenicity of Bd that is largely restricted to higher elevations and colder temperatures. These findings imply that the lowlands may play a critical role in the spread and persistence of Bd over time and space.

Studying the genetic factors underlying phenotypic traits can provide insight into dynamics of selection and molecular basis of adaptation, but this goal can be difficult for non-model organisms without extensive genomic resources. However, sequencing candidate genes for the trait of interest can facilitate the study of evolutionary genetics in natural populations. We sequenced the melanocortin-1 receptor (Mc1r) to study the genetic basis of color polymorphism in a group of snake species with variable black banding, the genera Sonora, Chilomeniscus, and Chionactis. Mc1r is an important gene in the melanin synthesis pathway and is associated with ecologically important variation in color pattern in birds, mammals, and other squamate reptiles. We found that Mc1r nucleotide sequence was variable and that within our focal Sonora species, there are both fixed and heterozygous nucleotide substitutions that result in an amino acid change and selection analyses indicated that Mc1r sequence was likely under purifying selection. However, we did not detect any statistical association with the presence or absence of black bands. Our results agree with other studies that have found no role for sequence variation in Mc1r and highlight the importance of comparative data for studying the phenotypic associations of candidate genes.

Batesian mimicry, in which harmless species (mimics) deter predators by deceitfully imitating the warning signals of noxious species (models), generates striking cases of phenotypic convergence that are classic examples of evolution by natural selection. However, mimicry of venomous coral snakes has remained controversial because of unresolved conflict between the predictions of mimicry theory and empirical patterns in the distribution and abundance of snakes. Here we integrate distributional, phenotypic and phylogenetic data across all New World snake species to demonstrate that shifts to mimetic coloration in nonvenomous snakes are highly correlated with coral snakes in both space and time, providing overwhelming support for Batesian mimicry. We also find that bidirectional transitions between mimetic and cryptic coloration are unexpectedly frequent over both long- and short-time scales, challenging traditional views of mimicry as a stable evolutionary ‘end point' and suggesting that insect and snake mimicry may have different evolutionary dynamics., Toxic and venomous species often have conspicuous warning colouration that is mimicked by harmless species. Here, Davis Rabosky et al. combine phylogenetic and biogeographic analyses to reveal that mimicry of venomous coral snakes has been a major driver of snake colour evolution in the New World.

The restricted distribution and isolation of island endemics often produces unique genetic and phenotypic diversity of conservation interest to management agencies. However, these isolated species, especially those with sensitive life history traits, are at high risk for the adverse effects of genetic drift and habitat degradation by non-native wildlife. Here, we study the population genetic diversity, structure, and stability of a classic “island giant” (Xantusia riversiana, the Island Night Lizard) on San Clemente Island, California following the removal of feral goats. Using DNA microsatellites, we found that this population is reasonably genetically robust despite historical grazing, with similar effective population sizes and genetic diversity metrics across all sampling locations irrespective of habitat type and degree of degradation. However, we also found strong site-specific patterns of genetic variation and low genetic diversity compared to mainland congeners, warranting continued special management as an island endemic. We identify both high and low elevation areas that remain valuable repositories of genetic diversity and provide a case study for other low-dispersal coastal organisms in the face of future climate change.

Mexico possesses high levels of endemic biodiversity, especially for squamate reptiles. However, the evolutionary relationships among many reptiles in this region are not well known. The closely related genera of Sonora Baird and Girard 1853 and Procinura Cope 1879 are coralsnake mimics found from the central and western United States to southwestern Mexico and Baja California. Although species delimitation in this group has historically relied upon colour pattern and other morphological characters, many populations of these species display colour pattern polymorphism, which may confound taxonomy. We used molecular phylogenetics to assess the evolutionary relationships and delimit species within Sonora, focusing on the phylogenetic position of Procinura and the validity of S. mutabilis and aequalis. We sequenced two mitochondrial (ND4 and cytb) and two nuclear (c-mos and RAG-1) genes for the single species of Procinura and each of the four species of Sonora. We analysed these sequences using maximum parsi...

Identifying the genetic basis of mimetic signals is critical to understanding both the origin and dynamics of mimicry over time. For species not amenable to large laboratory breeding studies, widespread color polymorphism across natural populations offers a powerful way to assess the relative likelihood of different genetic systems given observed phenotypic frequencies. We classified color phenotype for 2175 ground snakes (Sonora semiannulata) across the continental United States to analyze morph ratios and test among competing hypotheses about the genetic architecture underlying red and black coloration in coral snake mimics. We found strong support for a two-locus model under simple Mendelian inheritance, with red and black pigmentation being controlled by separate loci. We found no evidence of either linkage disequilibrium between loci or sex linkage. In contrast to Batesian mimicry systems such as butterflies in which all color signal components are linked into a single "supergene," our results suggest that the mimetic signal in colubrid snakes can be disrupted through simple recombination and that color evolution is likely to involve discrete gains and losses of each signal component. Both outcomes are likely to contribute to the exponential increase in rates of color evolution seen in snake mimicry systems over insect systems.

This report summarizes the proceedings of the 1st Snake Genomics and Integrative Biology Meeting held in Vail, CO USA, 5–8 October 2011. The meeting had over twenty registered participants, and was conducted as a single session of presentations. Goals of the meeting included coordination of genomic data collection and fostering collaborative interactions among researchers using snakes as model systems.

Color polymorphism in natural populations presents an ideal opportunity to study the evolutionary drivers of phenotypic diversity. Systems with striking spatial, temporal, and qualitative variation in color can be leveraged to study the mechanisms promoting the distribution of different types of variation in nature. We used the highly polymorphic ground snake (Sonora semiannulata), a putative coral snake mimic with both cryptic and conspicuous morphs, to compare patterns of neutral genetic variation and variation over space and time in color polymorphism to investigate the mechanistic drivers of phenotypic variation across scales. We found that strong selection promotes color polymorphism across spatial and temporal scales, with morph frequencies differing markedly between juvenile and adult age classes within a single population, oscillating over time within multiple populations, and varying drastically over the landscape despite minimal population genetic structure. However, we found no evidence that conspicuousness of morphs was related to which color pattern was favored by selection or to any geographic factors, including sympatry with coral snakes. We suggest that complex patterns of phenotypic variation in polymorphic systems may be a fundamental outcome of the conspicuousness of morphs and that explicit tests of temporal and geographic variation are critical to the interpretation of conspicuousness and mimicry.

Social aggregation is a common behavioral phenomenon thought to evolve through adaptive benefits to group living. Comparing fitness differences between aggregated and solitary individuals in nature – necessary to infer an evolutionary benefit to living in groups – has proven difficult because communally-living species tend to be obligately social and behaviorally complex. However, these differences and the mechanisms driving them are critical to understanding how solitary individuals transition to group living, as well as how and why nascent social systems change over time. Here we demonstrate that facultative aggregation in a reptile (the Desert Night Lizard, Xantusia vigilis) confers direct reproductive success and survival advantages and that thermal benefits of winter huddling disproportionately benefit small juveniles, which can favor delayed dispersal of offspring and the formation of kin groups. Using climate projection models, however, we estimate that future aggregation in night lizards could decline more than 50% due to warmer temperatures. Our results support the theory that transitions to group living arise from direct benefits to social individuals and offer a clear mechanism for the origin of kin groups through juvenile philopatry. The temperature dependence of aggregation in this and other taxa suggests that environmental variation may be a powerful but underappreciated force in the rapid transition between social and solitary behavior.