Sceloporus Research Articles

Exploring the causes underlying the latitudinal variation in range sizes: Evidence for Rapoport's rule in spiny lizards (genus Sceloporus).

Species' range size is a fundamental unit of analysis in biodiversity research, given its association with extinction risk and species richness. One of its most notable patterns is its positive relationship with latitude, which has been considered an ecogeographical rule called Rapoport's rule. Despite this rule being confirmed for various taxonomic groups, its validity has been widely discussed and several taxa still lack a formal assessment. Different hypotheses have been proposed to explain their potential mechanisms, with those related to temperature and elevational being the most supported thus far. In this study, we employed two level of analyses (cross-species and assemblage) to investigate the validity of Rapoport's rule in spiny lizards (genus Sceloporus). Additionally, we evaluated four environmental-related hypotheses (minimum temperature, temperature variability, temperature stability since the last glacial maximum, and elevation) posed to explain such pattern, contrasting our results to those patterns expected under a null model of range position. Our results provided support for Rapoport's rule at both levels of analyses, contrasting with null expectations. Consistently, minimum temperature and elevation were the most relevant variables explaining the spatial variation in range size. At the cross-species level, our null simulations revealed that both variables deviated significantly from random expectations. Conversely, at the assemblage level, none of the variables were statistically different from the expected relationships. We discussed the implication of our findings in relation to the ecology and evolution of spiny lizards.

Flexibility in thermal requirements: a comparative analysis of the wide-spread lizard genus Sceloporus.

Adaptation or acclimation of thermal requirements to environmental conditions can reduce thermoregulation costs and increase fitness, especially in ectotherms, which rely heavily on environmental temperatures for thermoregulation. Insight into how thermal niches have shaped thermal requirements across evolutionary history may help predict the survival of species during climate change. The lizard genus Sceloporus has a widespread distribution and inhabits an ample variety of habitats. We evaluated the effects of geographical gradients (i.e. elevation and latitude) and local environmental temperatures on thermal requirements (i.e. preferred body temperature, active body temperature in the field, and critical thermal limits) of Sceloporus species using published and field-collected data and performing phylogenetic comparative analyses. To contrast macro- and micro-evolutional patterns, we also performed intra-specific analyses when sufficient reports existed for a species. We found that preferred body temperature increased with elevation, whereas body temperature in the field decreased with elevation and increased with local environmental temperatures. Critical thermal limits were not related to the geographic gradient or environmental temperatures. The apparent lack of relation of thermal requirements to geographic gradient may increase vulnerability to extinction due to climate change. However, local and temporal variations in thermal landscape determine thermoregulation opportunities and may not be well represented by geographic gradient and mean environmental temperatures. Results showed that Sceloporus lizards are excellent thermoregulators, have wide thermal tolerance ranges, and the preferred temperature was labile. Our results suggest that Sceloporus lizards can adjust to different thermal landscapes, highlighting opportunities for continuous survival in changing thermal environments.

Gene Flow and Isolation in the Arid Nearctic Revealed by Genomic Analyses of Desert Spiny Lizards.

The opposing forces of gene flow and isolation are two major processes shaping genetic diversity. Understanding how these vary across space and time is necessary to identify the environmental features that promote diversification. The detection of considerable geographic structure in taxa from the arid Nearctic has prompted research into the drivers of isolation in the region. Several geographic features have been proposed as barriers to gene flow, including the Colorado River, Western Continental Divide (WCD), and a hypothetical Mid-Peninsular Seaway in Baja California. However, recent studies suggest that the role of barriers in genetic differentiation may have been overestimated when compared to other mechanisms of divergence. In this study, we infer historical and spatial patterns of connectivity and isolation in Desert Spiny Lizards (Sceloporus magister) and Baja Spiny Lizards (Sceloporus zosteromus), which together form a species complex composed of parapatric lineages with wide distributions in arid western North America. Our analyses incorporate mitochondrial sequences, genomic-scale data, and past and present climatic data to evaluate the nature and strength of barriers to gene flow in the region. Our approach relies on estimates of migration under the multispecies coalescent to understand the history of lineage divergence in the face of gene flow. Results show that the S. magister complex is geographically structured, but we also detect instances of gene flow. The WCD is a strong barrier to gene flow, while the Colorado River is more permeable. Analyses yield conflicting results for the catalyst of differentiation of peninsular lineages in S. zosteromus. Our study shows how large-scale genomic data for thoroughly sampled species can shed new light on biogeography. Furthermore, our approach highlights the need for the combined analysis of multiple sources of evidence to adequately characterize the drivers of divergence.

More cryptic diversity among spiny lizards of the Sceloporus torquatus complex discovered through a multilocus approach

Abstract A phylogenetic reassessment of the Sceloporus torquatus species complex is presented using a multilocus approach. Topological conflicts related to a clade belonging to disjunct southern populations of S. madrensis are resolved, confirming the identity of such populations as distinct from the nominal populations of S. madrensis located 175 kilometres to the north. We describe this southern clade as a new species distributed in the Sierra Madre Oriental based on its phylogenetic distinctiveness and a combination of morphological characters and measurements, increasing the number of recognised species in the S. torquatus complex to seven.

A NEW SPECIES OF CHIGGER MITE AND THE RECORD OF A GASTROINTESTINAL NEMATODE IN SCELOPORUS CAUTUS (SQUAMATA: PHRYNOSOMATIDAE) FROM THE CHIHUAHUAN DESERT PROVINCE IN COAHUILA, MEXICO.

The chigger species Acomatacarus coahuilensis n. sp. (Acariformes: Trombidiformes: Leeuwenhoekiidae) inhabiting the neck scales of the wary spiny lizard Sceloporus cautus Smith, 1938 (Squamata: Iguania: Phrynosomatidae), from Chihuahuan Desert Province, Coahuila, Mexico, is described herein. The new mite species is assigned to the genus AcomatacarusEwing, 1942, based on a palptarsus with 6 branched setae plus the tarsala (ω). This new species can be separated from the other species in this genus mainly based on the combination of the following characters: presence of mastisetae (MTa) on tarsus III and tibiala (φ) on tibia III, subterminala (ζ1), parasubterminala (z), and pretarsala (ζ2) on tarsus I, 2 genualae (σ1-2) on genu I, onychotriches and claw-like empodium of legs I-III, and absence of mastisetae on tibia III. In addition to the new species, the gastrointestinal nematode Skrjabinoptera phrynosoma (Ortlepp, 1922) Schulz, 1927 (Physalopteridae), was also found, which represents a new host record in the wary spiny lizard. This study reports the third species of the genus Acomatacarus recorded from Mexico and the first record of chigger mites and helminth parasites associated with Sc. cautus.

Population differences in multimodal lizard communication are not well explained by habitat or history

AbstractAnimals evolve in complex selective regimes, where a suite of different factors can shape signal use. We might predict that more closely related species will exhibit more similar behavior than those more distantly related; however, sometimes signals are shaped more profoundly by the environment or other forces. Lizards in the genus Sceloporus communicate with conspecifics with multimodal signals that combine species‐typical push‐up and headbob displays and chemical signals in the form of femoral gland secretions. Here, we examine behavioral activity and signal use across three closely related populations of the Sceloporus undulatus species complex from diverse habitats across the United States, to test the relative roles of habitat and phylogeography in shaping communicative behavior. We filmed undisturbed levels of activity for free‐ranging males of S. consobrinus, syn. S. u. erythrocheilus, in Colorado, S. undulatus hyacinthinus in Indiana and S. u. undulatus in Georgia, and scored frequency and rates of behavior important for communication. We found that populations differed in their use of communicative signals in a way that deviates from expectations based solely on phylogeographic proximity or habitat, suggesting that plasticity or adaptation to conditions that vary among populations may be especially important. Specifically, canonical discriminant analyses found the largest differences in movement patterns. Sceloporus u. hyacinthinus was the most behaviorally different out of the three: males in this population had lower movement rates and particularly low levels of chemosensory behavior while male S. consobrinus and S. u. undulatus showed similar rates of chemosensory acts and headbob/push‐up displays. Phenotypic and environmental variation among closely related populations, in combination with phylogeographic knowledge can help us untangle the processes responsible for the origin and maintenance of organismal diversity in communicative behavior.

Phenotypic diversity facilitates niche partitioning in a sky island assemblage of spiny lizards

Abstract If closely related species should be more similar and therefore more likely to compete for resources, why do we see so many examples of species-rich congeneric communities in nature? To avoid competition, many species possess suites of traits associated with differential resource use – or ecomorphologies – that promote niche differentiation with co-occurring species. Yet, the axes through which niche partitioning evolves and the traits involved are still poorly understood in most systems. Island systems, in which species interactions can form strong forces of selection due to limited resources, provide valuable insights into how ecomorphological diversity contributes to coexistence. Here, we examined axes of resource use and morphological traits that facilitate niche partitioning in a community of spiny lizards (genus Sceloporus) in a sky island system, the Chiricahua Mountains of southeastern Arizona, USA. By quantifying structural, temporal and thermal niche use for over 300 co-occurring lizards from four species over three field seasons, we show that sympatric species diverge primarily in perch height and type, but also in thermal and temporal resource use. Our results further demonstrate interspecific divergence in a suite of phenotypic traits known to covary with ecology in other lizard radiations, including body size, scale size, hindlimb length, foot length and dorsal coloration. Studies of niche partitioning and ecomorphology, especially of closely related species, deepen our understanding of how diverse communities assemble and how morphological diversity accumulates across the tree of life, and our findings highlight the importance of examining multiple axes of resource use to better understand these processes.

DNA metabarcoding reveals seasonal changes in diet composition across four arthropod-eating lizard species (Phrynosomatidae: Sceloporus).

Diet composition and its ecological drivers are rarely investigated in coexisting closely related species. We used a molecular approach to characterize the seasonal variation in diet composition in four spiny lizard species inhabiting a mountainous ecosystem. DNA metabarcoding revealed that the lizards Sceloporus aeneus, S. bicanthalis, S. grammicus, and S. spinosus mostly consumed arthropods of the orders Hemiptera, Araneae, Hymenoptera, and Coleoptera. The terrestrial lizards S. aeneus and S. bicanthalis mostly predated ants and spiders, whereas the arboreal-saxicolous S. grammicus and saxicolous S. spinosus largely consumed grasshoppers and leafhoppers. The taxonomic and phylogenetic diversity of the prey was higher during the dry season than the rainy season, likely because reduced prey availability in the dry season forced lizards to diversify their diets to meet their nutritional demands. Dietary and phylogenetic composition varied seasonally depending on the species, but only dietary composition varied with altitude. Seasonal dietary turnover was greater in S. spinosus than in S. bicanthalis, suggesting site-specific seasonal variability in prey availability; no other differences among species were observed. S. bicanthalis, which lives at the highest altitude in our study site, displayed interseasonal variation in diet breadth. Dietary differences were correlated with the species' feeding strategies and elevational distribution, which likely contributed to the coexistence of these lizard species in the studied geographic area and beyond.

Assembly of the largest squamate reference genome to date: The western fence lizard, Sceloporus occidentalis.

Spiny lizards (genus Sceloporus) have long served as important systems for studies of behavior, thermal physiology, dietary ecology, vector biology, speciation, and biogeography. The western fence lizard, Sceloporus occidentalis, is found across most of the major biogeographical regions in the western United States and northern Baja California, Mexico, inhabiting a wide range of habitats, from grassland to chaparral to open woodlands. As small ectotherms, Sceloporus lizards are particularly vulnerable to climate change, and S. occidentalis has also become an important system for studying the impacts of land use change and urbanization on small vertebrates. Here, we report a new reference genome assembly for S. occidentalis, as part of the California Conservation Genomics Project (CCGP). Consistent with the reference genomics strategy of the CCGP, we used Pacific Biosciences HiFi long reads and Hi-C chromatin-proximity sequencing technology to produce a de novo assembled genome. The assembly comprises a total of 608 scaffolds spanning 2,856 Mb, has a contig N50 of 18.9 Mb, a scaffold N50 of 98.4 Mb, and BUSCO completeness score of 98.1% based on the tetrapod gene set. This reference genome will be valuable for understanding ecological and evolutionary dynamics in S. occidentalis, the species status of the California endemic island fence lizard (S. becki), and the spectacular radiation of Sceloporus lizards.

Missing toes in the graphic spiny lizard Sceloporus grammicus from central Mexico

Missing toes in the graphic spiny lizard Sceloporus grammicus from central Mexico

Comparative analysis of two nonlethal methods for the study of the gut bacterial communities in wild lizards.

Fecal samples or cloacal swabs are preferred over lethal dissections to study vertebrate gut microbiota for ethical reasons, but it remains unclear which nonlethal methods provide more accurate information about gut microbiota. We compared the bacterial communities of three gastrointestinal tract (GIT) segments, that is, stomach, small intestine (midgut), and rectum (hindgut) with the bacterial communities of the cloaca and feces in the mesquite lizard Sceloporus grammicus. The hindgut had the highest taxonomic and functional alpha diversity, followed by midgut and feces, whereas the stomach and cloaca showed the lowest diversities. The taxonomic assemblages of the GIT segments at the phylum level were strongly correlated with those retrieved from feces and cloacal swabs (rs > 0.84 in all cases). The turnover ratio of Amplicon Sequence Variants (ASVs) between midgut and hindgut and the feces was lower than the ratio between these segments and the cloaca. More than half of the core-ASVs in the midgut (24 of 32) and hindgut (58 of 97) were also found in feces, while less than 5 were found in the cloaca. At the ASVs level, however, the structure of the bacterial communities of the midgut and hindgut were similar to those detected in feces and cloaca. Our findings suggest that fecal samples and cloacal swabs of spiny lizards provide a good approximation of the taxonomic assemblages and beta diversity of midgut and hindgut microbiota, while feces better represent the bacterial communities of the intestinal segments at a single nucleotide variation level than cloacal swabs.

UPPER ELEVATIONAL RANGE EXTENSION OF Sceloporus utiformis COPE, 1864 (SQUAMATA: PHRYNOSOMATIDAE) IN THE SIERRA DE MANANTLÁN BIOSPHERE RESERVE, MEXICO

Cope's Largescale Spiny Lizards (Sceloporus utiformis) are endemic to the Pacific slope of Mexico. They are oviparous and are known to inhabit tropical and temperate forests from sea level to 1,800 m a.s.l. Here, we expand this range based on the finding, in May 2022, of two individuals of S. utiformis in the southeastern portion of the Sierra de Manantlán, at 2154 m elevation. We also provide data on morphometry and field body and microhabitat temperatures of these organisms. A previous study conducted at the same locality almost three decades ago found the species to be present at much lower elevations, so we hypothesize that this could be a case of upward range shifting due to climate change. Moreover, as few oviparous members of the genus Sceloporus occur above 2,000 m elevation, this record triggers many questions about the ecological implications of oviparity along elevation gradients.

Cloacal microbiomes of sympatric and allopatric Sceloporus lizards vary with environment and host relatedness.

Animals and their microbiomes exert reciprocal influence; the host's environment, physiology, and phylogeny can impact the composition of the microbiome, while the microbes present can affect host behavior, health, and fitness. While some microbiomes are highly malleable, specialized microbiomes that provide important functions can be more robust to environmental perturbations. Recent evidence suggests Sceloporus virgatus has one such specialized microbiome, which functions to protect eggs from fungal pathogens during incubation. Here, we examine the cloacal microbiome of three different Sceloporus species (spiny lizards; Family Phrynosomatidae)-Sceloporus virgatus, Sceloporus jarrovii, and Sceloporus occidentalis. We compare two species with different reproductive modes (oviparous vs. viviparous) living in sympatry: S. virgatus and S. jarrovii. We compare sister species living in similar habitats (riparian oak-pine woodlands) but different latitudes: S. virgatus and S. occidentalis. And, we compare three populations of one species (S. occidentalis) living in different habitat types: beach, low elevation forest, and the riparian woodland. We found differences in beta diversity metrics between all three comparisons, although those differences were more extreme between animals in different environments, even though those populations were more closely related. Similarly, alpha diversity varied among the S. occidentalis populations and between S. occidentalis and S. virgatus, but not between sympatric S. virgatus and S. jarrovii. Despite these differences, all three species and all three populations of S. occcidentalis had the same dominant taxon, Enterobacteriaceae. The majority of the variation between groups was in low abundance taxa and at the ASV level; these taxa are responsive to habitat differences, geographic distance, and host relatedness. Uncovering what factors influence the composition of wild microbiomes is important to understanding the ecology and evolution of the host animals, and can lead to more detailed exploration of the function of particular microbes and the community as a whole.

NEW HIGHEST ELEVATION RECORDS OF TWO MEXICAN ENDEMIC SPINY LIZARDS: Sceloporus megAlepidurus AND Sceloporus spinosus (SQUAMATA: PHRYNOSOMATIDAE)

We record the new highest elevation of two endemic lizards from Mexico. Our records for the Largescale Spiny Lizard Sceloporus megalepidurus Smith, 1934 extends 310 m its previous highest record, from 3,080 to 3,390 m in Peñon de Tlaxco, and for the Eastern Spiny Lizard S. spinosus Wiegmann, 1828 extends 360 m its previous highest record, from 2,670 to 3,030 m in the volcano La Malinche, both records in Tlaxcala, Mexico. We discuss our observations in the context of global climate change, habitat alterations, and the ecological implications in the new environment.

An annotated list of the species-group names applied to the lizard genus Sceloporus

An alphabetically-arranged list of all (196) species-group names ever proposed or used in the genus Sceloporus is presented, with citations of the original proposal, onomatophore, type locality, English standardnames, current status, combination priority, group assignment, and etymology. Twenty-three species groups, 91 species (66 monotypic) and 153 species- group names are recognized as valid.

A new species of Sceloporus of the torquatus group (Reptilia: Phrynosomatidae) from West Mexico.

We describe Sceloporus huichol sp. nov., a new species of spiny lizard of the genus Sceloporus, that is found in the mountainous regions of Jalisco and Nayarit in western Mexico. The new species belongs to the torquatus species group and has previously been confused with Sceloporus melanogaster, from which it differs by its smaller size, head color and nuchal collar. Additionally, Sceloporus huichol sp. nov. is different from the rest of its congeners by relevant genetic and morphologicall data. Despite having a complete nuchal collar, characteristic of Sceloporus torquatus, S. huichol sp. nov. is more closely related phylogenetically to S. melanogaster. Recognition of this new species brings the number of species in the torquatus group to nine.

08. Red tail in male spiny footed lizards Acanthodactylus erythrurus during the breeding season

08. Red tail in male spiny footed lizards Acanthodactylus erythrurus during the breeding season

Red tail in male spiny footed lizards Acanthodactylus erythrurus during the breeding season

Red tail in male spiny footed lizards Acanthodactylus erythrurus during the breeding season

A new and very spiny lizard (Gymnophthalmidae: Echinosaura) from the Andes in northwestern Ecuador.

We describe a new species of Neotropical spiny-lizard of the genus Echinosaura from the Imbabura and Carchi Provinces on the western slopes of the Andes in northwestern Ecuador. The new species mostly resembles E. horrida. However, it can be distinguished from all congeners by having keeled enlarged dorsal scales forming a paired vertebral row, two paravertebral series of short oblique rows of projecting scales, and a pair of spine-like scales on temporal and nuchal regions. We also provide a detailed description of the osteology of the skull and pectoral girdle of the new species and present a phylogenetic hypothesis for Echinosaura based on three mitochondrial genes (12S, 16S, ND4) and one nuclear gene (c-mos).

Hidden diversity within a polytypic species: The enigmatic Sceloporus torquatus Wiegmann, 1828 (Reptilia, Squamata, Phrynosomatidae)

The spiny lizard genus Sceloporus was described by Wiegmann in 1828, with S. torquatus posteriorly designated as the type species. The taxonomic history of S. torquatus is complicated, as it has been confused with other taxa by numerous authors. Many modern systematics works have been published on Sceloporus, but none have included all five recognized S. torquatus subspecies: S. t. torquatus, S. t. melanogaster, S. t. binocularis, S. t. mikeprestoni, and S. t. madrensis. Additionally, there is previous evidence for at least one unnamed taxon. The present study is the first taxonomic revision of the enigmatic S. torquatus based on molecular phylogenies using combined molecular data from 12S, ND4 and RAG1 genes, and Maximum Likelihood and Bayesian inference phylogenetic methods. This work includes the most extensive sampling across the entire distribution, as well as divergence time estimates and environmental niche modelling, which combined offer a spatio-temporal framework for understanding the evolution of the species. Additionally, a series of morphological characters are analyzed to identify significant differences between lineages consistently recovered in the molecular phylogenies. Using this integrative approach, evidence is presented for eight lineages within the S. torquatus complex, five of which correspond to previously recognized subspecies and three represent unnamed taxa masked by morphological conservatism. Finally, to maintain taxonomic stability a lectotype and paralectoype are designated for S. torquatus, and certain taxonomic changes are suggested in order to reflect the phylogenetic relationships within the S. torquatus complex.