Geology topics
Publications and source records attributed to Taylor Edwards.
Sonoran Desert Tortoise Gopherus morafkai has most recently been assessed for The IUCN Red List of Threatened Species in 2019. Gopherus morafkai is listed as Vulnerable under criteria A2abce+4abce.
Sampling fecal droppings (scat) to genetically identify individual animals is an established method for monitoring mammal populations and could be highly useful for monitoring reptile populations. Whereas existing protocols for obtaining DNA from reptile scat focus on analyses of whole, fresh scat deposited during animal handling, the collection of scat naturally deposited by reptiles in situ, as required for non-invasive population monitoring, requires protocols to extract highly degraded DNA. Using surface swabs from such scats can reduce PCR inhibition, ecological impacts of removing scat, and zoonotic risks. We report on three related but independently designed studies of DNA analyses from scat swabs of herbivorous reptiles under natural desert conditions: two free-ranging desert tortoise species (Agassiz's desert tortoise, Gopherus agassizii , California, US, and Morafka's desert tortoise, G. morafkai , Arizona, US) and the common chuckwalla (Sauromalus atar ) (Arizona, US, and Sonora, MX). We analyzed samples from both tortoise species with the same set of 16 microsatellites and chuckwalla samples with four mtDNA markers; studies also varied in swab preservation medium and DNA extraction method. Microsatellite amplification success per sample, defined as ≥9 loci with amplification, varied between studies, with 15% for Agassiz's desert tortoise and 42% Morafka's desert tortoise. For chuckwallas, we successfully amplified and sequenced 50% of samples. We recovered fragments up to 400 bp for tortoises and 980 bp for chuckwallas from scat swab samples. This study demonstrates that genotypes can successfully be obtained from swabs of herbivorous reptile scat collected in the field under natural environmental conditions and emphasizes that repeat amplifications are necessary for genetic identification of individuals from non-invasive samples.
Understanding the influence of geographic features on the evolutionary history and population structure of a species can assist wildlife managers in delimiting genetic units (GUs) for conservation and management. Landscape features including mountains, low elevation depressions, and even roads can influence connectivity and gene flow among Agassiz’s desert tortoise (Gopherus agassizii) populations. Substantial changes in the landscape of the American Southwest occurred during the last six million years (including the formation of the Gulf of California and the lower Colorado River), which shaped the distribution and genetic structuring of tortoise populations. The area northwest of the Gulf of California is occupied by the Salton Trough, including the Coachella Valley at its northern end. Much of this area is below sea level and unsuitable as tortoise habitat, thus forming a potential barrier for gene flow. We assessed genetic relationships among three tortoise populations separated by the Coachella Valley. Two adjacent populations were on the east side of the valley in the foothills of the Cottonwood and Orocopia mountains separated by Interstate 10. The third population, Mesa, was located about 87 km away in the foothills of the San Bernardino Mountains at the far northwestern tip of the valley. The Cottonwood and Orocopia localities showed genetic affiliation with the adjacent Colorado Desert GU immediately to the east, and the Mesa population exhibited affiliation with both the Southern Mojave and Colorado Desert GUs, despite having a greater geographic distance (0.5x–1.5x greater) to the Colorado Desert GU. The genetic affiliation with the Colorado Desert GU suggests that the boundary for that GU needs to be substantially extended to the west to include the desert tortoise populations around the Coachella Valley. Their inclusion in the Colorado Desert GU may benefit these often overlooked populations when recovery actions are considered.
Despite an historical connection between the arts and sciences, in the past century, the two disciplines have been greatly siloed. However, there is a renewed interest in collaboration across the arts and sciences to support conservation practice by understanding and communicating complex environmental, social, and cultural challenges in novel ways. 6&6 was created as a transdisciplinary art–science initiative to promote a deeper appreciation of the Sonoran Desert. Six artists and six scientists were paired to create work that explored conservation issues in the Sonoran Desert and the Gulf of California. In-depth interviews were conducted with the artists and scientists throughout the 4-year initiative to understand the impact of 6&6 on their personal and professional behaviors and outlook. The findings from this case study reveal the role that intensive, place-based, and transdisciplinary art–science programs can play in shaping narratives to better communicate the patterns and processes of nature and human–environment interactions.
Turtles and tortoises (chelonians) have been integral components of global ecosystems for about 220 million years and have played important roles in human culture for at least 400,000 years. The chelonian shell is a remarkable evolutionary adaptation, facilitating success in terrestrial, freshwater and marine ecosystems. Today, more than half of the 360 living species and 482 total taxa (species and subspecies combined) are threatened with extinction. This places chelonians among the groups with the highest extinction risk of any sizeable vertebrate group. Turtle populations are declining rapidly due to habitat loss, consumption by humans for food and traditional medicines and collection for the international pet trade. Many taxa could become extinct in this century. Here, we examine survival threats to turtles and tortoises and discuss the interventions that will be needed to prevent widespread extinction in this group in coming decades.
Aims To investigate spatial congruence between ecological niches and genotype in two allopatric species of desert tortoise that are species of conservation concern. Location Mojave and Sonoran Desert ecoregions; California, Nevada, Arizona, Utah, USA. Methods We compare ecological niches of Gopherus agassizii and Gopherus morafkai using species distribution modelling (SDM) and then calibrate a pooled‐taxa distribution model to explore local differences in species–environment relationships based on the spatial residuals of the pooled‐taxa model. We use multiscale geographically weighted regression (MGWR) applied to those residuals to estimate local species–environment relationships that can vary across the landscape. We identify multivariate clusters in these local species–environment relationships and compare them against models of (a) a geographically based taxonomic designation for two sister species and (b) an environmental ecoregion designation, with respect to their ability to predict a genotype association index for these two species. Results We find non‐identical niches for these species, with differences that span physiographic and vegetation niche dimensions. We find evidence for two distinct clusters of local species–environment relationships that when mapped, predict an index of genotype association for the two sister taxa better than did either the geographically based taxonomic designation or an environmental ecoregion designation. Main conclusions Exploring local species–environment relationships by coupling SDM and MGWR can benefit studies of biogeography and conservation. We find that niche separation in habitat selection conforms to genotypic differences between sister taxa of tortoise in a recent secondary contact zone. This result may inform decision making by agencies with regulatory or land management authority for the two sister taxa addressed here.
This report summarizes the results for mtDNA and STR genotyping of 41 desert tortoise (Gopherus agassizii) DNA samples from opposite sides of the Coachella Valley: one sample from the west side at the Mesa wind energy facility in the Whitewater Hills and the other from the mouth of Cottonwood Canyon in Joshua Tree National Park, both within the boundaries of the Coachella Valley Multiple Species Habitat Conservation Plan. Additional samples were collected from tortoises on the northern bajadas of the Orocopia Mountains and from the Santa Rosa Mountains and those results will be presented at a later time in a scientific publication. We tested samples for their mtDNA haplotype and 25 STR loci previously used in other studies. We performed assignment testing to determine the genetic affinity of each individual to the geographic region of collection. Despite apparent isolation, both populations appear to be naturally occurring and do exhibit indications of having experienced increased genetic drift (resulting in increased homozygosity, increased inbreeding or a reduction of genetic diversity). The lack of strong evidence for genetic isolation suggests that long-term maintenance of unfragmented landscapes is an important part of tortoise conservation in the region.
Evolutionary biology often seeks to decipher the drivers of speciation, and much debate persists over the relative importance of isolation and gene flow in the formation of new species. Genetic studies of closely related species can assess if gene flow was present during speciation, because signatures of past introgression often persist in the genome. We test hypotheses on which mechanisms of speciation drove diversity among three distinct lineages of desert tortoise in the genus Gopherus . These lineages offer a powerful system to study speciation, because different biogeographic patterns (physical vs. ecological segregation) are observed at opposing ends of their distributions. We use 82 samples collected from 38 sites, representing the entire species' distribution and generate sequence data for mtDNA and four nuclear loci. A multilocus phylogenetic analysis in *BEAST estimates the species tree. RNA‐seq data yield 20,126 synonymous variants from 7665 contigs from two individuals of each of the three lineages. Analyses of these data using the demographic inference package ∂a∂i serve to test the null hypothesis of no gene flow during divergence. The best‐fit demographic model for the three taxa is concordant with the *BEAST species tree, and the ∂a∂i analysis does not indicate gene flow among any of the three lineages during their divergence. These analyses suggest that divergence among the lineages occurred in the absence of gene flow and in this scenario the genetic signature of ecological isolation (parapatric model) cannot be differentiated from geographic isolation (allopatric model).
Aim We examine the role biogeographical features played in the evolution of Morafka's desert tortoise ( Gopherus morafkai ) and test the hypothesis that G. morafkai maintains genetically distinct lineages associated with different Sonoran Desert biomes. Increased knowledge of the past and present distribution of the Sonoran Desert region's biota provides insight into the forces that drive and maintain its biodiversity. Location Sonoran Desert biogeographical region; Sonora and Sinaloa, Mexico and Arizona, USA. Methods We examined wild tortoises from Mexico ( n = 155) and Arizona ( n = 78), spanning their known distribution. We used mtDNA sequences to reconstruct matrilineal relationships and 25 microsatellite (STR) loci for Bayesian analyses of gene flow. We performed clinal analyses on both mtDNA and STR loci to determine the position and amount of introgression where lineages co-occur. We used GIS to assess the association of genetic structuring with ecological features. We used these data in a hypothesis-driven approach to assess different models of how genetic diversity is maintained and distributed in G. morafkai . Results Gopherus morafkai was found to comprise genetically and geographically distinct ‘Sonoran’ and ‘Sinaloan’ lineages. Both lineages occurred in a relatively narrow zone of overlap in Sinaloan thornscrub, where it transitions into Sonoran desertscrub. Limited introgression occurred at the contact zone. The best-fit model suggests that these lineages diverged in parapatry where the distribution of genotypes is environment-dependent and introgression is inhibited by exogenous selection. Main conclusions The historically shifting ecotone between tropical deciduous forest and Sonoran desertscrub appears to be a boundary that fostered divergence between parapatric lineages of tortoises. The sharp genetic cline between the two lineages suggests that periods of isolation in temporary refugia due to Pleistocene climatic cycling influenced divergence. Despite incomplete reproductive isolation, the Sonoran and Sinaloan lineages of G. morafkai are on separate evolutionary trajectories.
We examined a secondary contact zone between two species of desert tortoise, Gopherus agassizii and G. morafkai . The taxa were isolated from a common ancestor during the formation of the Colorado River (4-8 mya) and are a classic example of allopatric speciation. However, an anomalous population of G. agassizii comes into secondary contact with G. morafkai east of the Colorado River in the Black Mountains of Arizona and provides an opportunity to examine reinforcement of species' boundaries under natural conditions. We sampled 234 tortoises representing G. agassizii in California ( n = 103), G. morafkai in Arizona ( n = 78), and 53 individuals of undetermined assignment in the contact zone including and surrounding the Black Mountains. We genotyped individuals for 25 STR loci and determined maternal lineage using mtDNA sequence data. We performed multilocus genetic clustering analyses and used multiple statistical methods to detect levels of hybridization. We tested hypotheses about habitat use between G. agassizii and G. morafkai in the region where they co-occur using habitat suitability models. Gopherus agassizii and G. morafkai maintain independent taxonomic identities likely due to ecological niche partitioning, and the maintenance of the hybrid zone is best described by a geographical selection gradient model.
The conservation of tortoises poses a unique situation because several threatened species are commonly kept as pets within their native ranges. Thus, there is potential for captive populations to be a reservoir for repatriation efforts. We assess the utility of captive populations of the threatened Agassiz’s desert tortoise ( Gopherus agassizii ) for recovery efforts based on genetic affinity to local areas. We collected samples from 130 captive desert tortoises from three desert communities: two in California (Ridgecrest and Joshua Tree) and the Desert Tortoise Conservation Center (Las Vegas) in Nevada. We tested all samples for 25 short tandem repeats and sequenced 1,109 bp of the mitochondrial genome. We compared captive genotypes to a database of 1,258 Gopherus samples, including 657 wild caught G. agassizii spanning the full range of the species. We conducted population assignment tests to determine the genetic origins of the captive individuals. For our total sample set, only 44 % of captive individuals were assigned to local populations based on genetic units derived from the reference database. One individual from Joshua Tree, California, was identified as being a Morafka’s desert tortoise, G. morafkai , a cryptic species which is not native to the Mojave Desert. Our data suggest that captive desert tortoises kept within the native range of G. agassizii cannot be presumed to have a genealogical affiliation to wild tortoises in their geographic proximity. Precautions should be taken before considering the release of captive tortoises into the wild as a management tool for recovery.
The enduring processes of time, climate, and adaptation have sculpted the distribution of organisms we observe in the Sonoran Desert. One such organism is Morafka’s desert tortoise, Gopherus morafkai. We apply a genomic approach to identify the evolutionary processes driving diversity in this species and present preliminary findings and emerging hypotheses. The Sonoran Desert form of the tortoise exhibits a continuum of genetic similarity spanning 850 km of Sonoran desertscrub extending from Empalme, Sonora, to Kingman, Arizona. However, at the ecotone between desertscrub and foothills thornscrub we identify a distinct, Sinaloan lineage and this occurrence suggests a more complex evolutionary history for G. morafkai. By using multiple loci from throughout the tortoise’s genome, we aim to determine if divergence between these lineages occurred in allopatry, and further to investigate for signatures of past or current genetic introgression. This international, collaborative project will assist state and federal agencies in developing management strategies that best preserve the evolutionary potential of Morafka’s desert tortoise. Ultimately, an understanding of the evolutionary history of desert tortoises will not only clarify the forces that have driven the divergence in this group, but also contribute to our knowledge of the biogeographic history of the Southwestern deserts and how diversity is maintained within them.
We investigate a cornucopia of problems associated with the identity of the desert tortoise, Gopherus agassizii Cooper. The date of publication is found to be 1861, rather than 1863. Only one of the three original cotypes exists, and it is designated as the lectotype of the species. Another cotype is found to have been destroyed in the 1906 San Francisco earthquake and subsequent fire. The third is lost. The lectotype is genetically confirmed to be from California, and not Arizona, USA as sometimes reported. Maternally, the holotype of G. lepidocephalus Ottley et Velázques Solis, 1989 from the Cape Region of Baja California Sur, Mexico is also from the Mojavian population of the desert tortoise, and not from Tiburon Island, Sonora, Mexico as previously proposed. A suite of characters serve to diagnose tortoises west and north of the Colorado River, the Mojavian population, from those east and south of the river in Arizona, USA and Sonora and Sinaloa, Mexico, the Sonoran population. Species recognition is warranted and because G. lepidocephalus is from the Mojavian population no names are available for the Sonoran species. Thus, a new species, Gopherus morafkai sp. n., is named and this action reduces the distribution of G. agassizii to only 30% of its former range. This reduction has important implications for the conservation and protection of G. agassizii , which may deserve a higher level of protection.