H. Bradley Shaffer

• University of California, Los Angeles

Premise Azolla is a genus of floating ferns that has closely evolved with a vertically transmitted obligate cyanobacterium endosymbiont—Anabaena azollae—that fixes nitrogen. There are also other lesser-known Azolla symbionts whose role and mode of transmission are unknown. Methods We sequenced 112 Azolla specimens collected across the state of C...

Genetic rescue, or the translocation of individuals among populations to augment gene flow, can help ameliorate inbreeding depression and loss of adaptive potential in small and isolated populations. Genetic rescue is currently under consideration for an endangered butterfly in Canada, the Half‐moon Hairstreak ( Satyrium semiluna ). A small, unique...

Aim Our study provides foundational resources for future SDMing: methods for generating fine-scale, equal-area predictor datasets and best-practice SDM guidelines. We also provide reproducible code to streamline their implementation. Location Southwestern North America Methods Using over 215,000 research-grade iNaturalist occurrence records for 127...

Climate change and infectious disease jointly impact species worldwide. In addition to causing conspicuous mortality events, these threats produce a range of non‐lethal effects that are often overlooked, yet can affect individual survival and fecundity, and ultimately, population and species viability. We develop an energetic framework that structu...

Genetic rescue, or the translocation of individuals among populations to augment gene flow, can help ameliorate inbreeding depression and loss of adaptive potential in small and isolated populations. Genetic rescue is currently under consideration for an endangered butterfly in Canada, the half-moon hairstreak (Satyrium semiluna). A small, unique p...

The winter ant, Prenolepis imparis, is one of the most common, widespread, and conspicuous ant species in North America. P. imparis is well adapted to cold climates, and consequently, is often noted as the only active ant species during colder months. This specialized life history makes P. imparis a useful model organism for exploring thermal physi...

Established invasive species represent one of the most harmful and challenging threats to native biodiversity, necessitating methods for Early Detection and Rapid Response. Cryptic invasions are particularly challenging and often require expensive and time-consuming molecular surveys which limits their usefulness for management. We present a novel...

Azolla is a genus of freshwater ferns that is economically important as a nitrogen-fixing biofertilizer, biofuel, bioremediator, and for potential carbon sequestration, but also contains weedy invasive species. In California, only two species are currently recognized but there may be up to six putative species, with the discrepancy being due to the...

Salamanders have large and complex genomes, hampering whole genome sequencing, but reduced representation sequencing provides a feasible alternative. We present NewtCap: a sequence capture bait set that targets c.7k coding regions across the genomes of all True Salamanders and Newts (the family Salamandridae, also known as "salamandrids"). We test...

The federally endangered sister species, Eucyclogobius newberryi (northern tidewater goby) and E. kristinae (southern tidewater goby) comprise the California endemic genus Eucyclogobius, which historically occurred in all coastal California counties. Isolated lagoons that only intermittently connect to the sea are their primary habitat. Reproductio...

The federally endangered sister species, Eucyclogobius newberryi (northern tidewater goby) and E. kristinae (southern tidewater goby) comprise the California endemic genus Eucyclogobius, which historically occurred in all coastal California counties. Isolated lagoons that only intermittently connect to the sea are their primary habitat. Reproductio...

Salamanders of the genus Lyciasalamandra are represented by as many as 20 narrow-range endemic taxa inhabiting the Mediterranean coast of Turkey and a handful of Aegean Islands. Despite recent molecular phylogenetic studies, the genus is rife with uncertainty about the number of contained species and their phylogenetic relationships, both of which...

We describe a highly contiguous and complete diploid genome assembly for the Chryxus Arctic, Oeneis chryxus (E. Doubleday, [1849]), a butterfly species complex spanning much of northern and western North America. One subspecies, the Ivallda Arctic (O. c. ivallda), is endemic to California’s Sierra Nevada and of particular biogeographic interest and...

Polyergus kidnapper ants are widely distributed, but relatively uncommon, throughout the Holarctic, spanning an elevational range from sea level to over 3,000 m. These species are well known for their obligate social parasitism with various Formica ant species, which they kidnap in dramatic, highly coordinated raids. Kidnapped Formica larvae and pu...

When introduced species hybridize with native relatives, spread of advantageous invasive genes into native populations (introgression) is a conservation concern. Genome-scale SNP (single nucleotide polymorphism) analysis can be a powerful approach to detect hybridization and identify candidate loci experiencing selection in hybrid zones. However, f...

Azolla is a floating fern that has closely evolved with a vertically transmitted obligate cyanobacterium endosymbiont--Anabaena azollae that performs nitrogen fixation in specialized Azolla leaf pockets. This cyanobacterium has a greatly reduced genome and appears to be in the "advanced" stages of symbiosis, potentially evolving into a nitrogen fix...

The California Pipevine, Aristolochia californica Torr., is the only endemic California species within the cosmopolitan birthwort family Aristolochiaceae. It occurs as an understory vine in riparian and chaparral areas and in forest edges and windrows. The geographic range of this plant species almost entirely overlaps with that of its major specia...

Single nucleotide polymorphism (SNP) genotyping has become the default strategy for genetic analyses of natural populations. However, because of their simplicity, SNPs can sometimes be misleading. We used a seemingly well-curated panel of diagnostic SNPs to evaluate patterns of hybridization between introduced and native tiger salamanders ( Ambysto...

Carpenter ants in the genus Camponotus are large, conspicuous ants that are abundant and ecologically influential in many terrestrial ecosystems. The bicolored carpenter ant, Camponotus vicinus Mayr, is distributed across a wide range of elevations and latitudes in western North America, where it is a prominent scavenger and predator. Here, we pres...

The Yuma myotis bat (Myotis yumanensis) is a small vespertilionid bat and one of 52 species of new world Myotis bats in the subgenus Pizonyx. While M. yumanensis populations currently appear relatively stable, it is one of twelve bat species known or suspected to be susceptible to white-nose syndrome, the fungal disease causing declines in bat popu...

Snakes in the family Colubridae include over 2,000 currently recognized species, and comprise roughly 75% of the global snake species diversity on Earth. For such a spectacular radiation, colubrid snakes remain poorly understood ecologically and genetically. Two subfamilies, Colubrinae (788 species) and Dipsadinae (833 species), comprise the bulk o...

Invasive species present a global threat to the conservation of biodiversity. When invasive and endangered native taxa hybridize, the resulting admixture introduces novel conservation challenges. Across a large region of central California, a hybrid swarm consisting of admixed endangered California tiger salamanders (CTS) and introduced barred tige...

Rattlesnakes play important roles in their ecosystems by regulating prey populations, are involved in complex coevolutionary dynamics with their prey, and exhibit a variety of unusual adaptations, including maternal care, heat-sensing pit organs, hinged fangs, and medically-significant venoms. The western rattlesnake (Crotalus oreganus) is one of t...

The California Pipevine Swallowtail Butterfly, Battus philenor hirsuta, and its host plant, the California Pipevine or Dutchman's Pipe, Aristolochia californica Torr., are an important California endemic species pair. While this species pair is an ideal system to study co-evolution, genomic resources for both are lacking. Here, we report a new, chr...

Carpenter ants in the genus Camponotus are large, conspicuous ants that are abundant and ecologically influential in many terrestrial ecosystems. The bicolored carpenter ant, C. vicinus Mayr, is distributed across a wide range of elevations and latitudes in western North America, where it is a prominent scavenger and predator. Here, we present a hi...

Wide-ranging species are seldom considered conservation priorities, yet they have the potential to harbour genetically deeply differentiated units across environments or ecological barriers, including some that warrant taxonomic recognition. Documenting such cryptic genetic diversity is especially important for wide-ranging species that are in decl...

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, Mex...

Introduction A major goal for conservation planning is the prioritized protection and management of areas that harbor maximal biodiversity. However, such spatial prioritization often suffers from limited data availability, resulting in decisions driven by a handful of iconic or endangered species, with uncertain benefits for co-occurring taxa. We a...

Damselflies and dragonflies (Order: Odonata) play important roles in both aquatic and terrestrial food webs and can serve as sentinels of ecosystem health and predictors of population trends in other taxa. The habitat requirements and limited dispersal of lotic damselflies make them especially sensitive to habitat loss and fragmentation. As such, l...

Genome assemblies are increasingly being used to identify adaptive genetic variation that can help prioritize the population management of protected species. This approach may be particularly relevant to species like Blainville's horned lizard, Phrynosoma blainvillii, due to its specialized diet on noxious harvester ants, numerous adaptative traits...

The black rail, Laterallus jamaicensis, is one of the most secretive and poorly understood birds in the Americas. Two of its five subspecies breed in North America: the Eastern black rail (L. j. jamaicensis), found primarily in the southern and mid-Atlantic states, and the California black rail (L. j. coturniculus), inhabiting California and Arizon...

The Virginia rail, Rallus limicola, is a member of the family Rallidae, which also includes many other species of secretive and poorly studied wetland birds. It is recognized as a single species throughout its broad distribution in North America where it is exploited as a game bird, often with generous harvest limits, despite a lack of systematic p...

Speciation entails a reduction in gene flow between lineages. The rates at which genomic regions become isolated varies across space and time. Barrier markers are linked to putative genes involved in (processes of) reproductive isolation, and, when observed over two transects, indicate species-wide processes. In contrast, transect-specific putative...

In Southern California, irrigation infrastructure is a prerequisite for urban green space management, and valve boxes are installed widely to manage water flow. These below-ground, plastic boxes protect valves and manifolds, create space for connecting pipes, and present a scarce ecological resource—elevated humidity and shelter from potential pred...

Climate-driven changes in hydrological regimes are of global importance and are particularly significant in riparian ecosystems. Riparian ecosystems in California provide refuge to many native and vulnerable species within a xeric landscape. California Tetragnatha spiders play a key role in riparian ecosystems, serving as a link between terrestrial...

Sea turtles represent an ancient lineage of marine vertebrates that evolved from terrestrial ancestors over 100 Mya. The genomic basis of the unique physiological and ecological traits enabling these species to thrive in diverse marine habitats remains largely unknown. Additionally, many populations have drastically declined due to anthropogenic ac...

Integral projection models (IPMs) can estimate the population dynamics of species for which both discrete life stages and continuous variables influence demographic rates. Stochastic IPMs for imperiled species, in turn, can facilitate population viability analyses (PVAs) to guide conservation decision‐making. Biphasic amphibians are globally distri...

We developed a species-specific, quantitative PCR assay multiplex for the detection of two threatened and endangered anuran species, arroyo toad (Anaxyrus californicus) and western spadefoot (Spea hammondii), from environmental DNA (eDNA). Both species have experienced dramatic declines over the last century throughout their ranges, mostly as a res...

Pricklebacks (Family Stichaeidae) are generally cold-temperate fishes most commonly found in the north Pacific. As part of the California Conservation Genomics Project (CCGP), we sequenced the genome of the Monkeyface Prickleback, Cebidichthys violaceus, to establish a genomic model for understanding phylogeographic patterns of marine organisms in...

Conservation science and environmental regulation are sibling constructs of the latter half of the 20th century, part of a more general awakening to humanity’s effect on the natural world in the wake of two world wars. Efforts to understand the evolution of biodiversity using the models of population genetics and the data derived from DNA sequencin...

Sculpins (Family Cottidae) are generally cold-temperate intertidal reef fishes most commonly found in the North Pacific. As part of the California Conservation Genomics Project (CCGP), we sequenced the genome of the Woolly Sculpin, Clinocottus analis, to establish a genomic model for understanding phylogeographic structure of inshore marine taxa al...

Linanthus parryae is a widespread annual plant species found in washes and sandy open habitats across the Mojave Desert and Eastern Sierra Nevada of California. Studies in this species have played a central role in evolutionary biology, serving as the first test cases of the shifting balance theory of evolution, models of isolation by distance, and...

A recently published macrogenetic dataset of California’s flora and fauna, CaliPopGen, comprehensively summarizes population genetic research published between 1985-2020. Integrating these genetic data into the requisite “best available science” upon which conservation professionals rely should facilitate the prioritization of populations based on...

The rubber boa, Charina bottae is a semi-fossorial, cold-temperature adapted snake that ranges across the wetter and cooler ecoregions of the California Floristic Province. The rubber boa is one of two species in the family Boidae native to California and currently has two recognized subspecies, the Northern rubber boa Charina b. bottae and the Sou...

Incorporating measures of taxonomic diversity into research and management plans has long been a tenet of conservation science. Increasingly, active conservation programs are turning towards multi-species landscape and regional conservation actions, and away from single species approaches. This is both a reflection of changing trends in conservatio...

The glossy snake (Arizona elegans) is a polytypic species broadly distributed across southwestern North America. The species occupies habitats ranging from California’s coastal chaparral to the shortgrass prairies of Texas and southeastern Nebraska, to the extensive arid scrublands of central México. Three subspecies are currently recognized in Cal...

Surfperches (Family Embiotocidae) are viviparous temperate reef fishes that brood their young. This life history trait translates into limited dispersal, strong population structure, and an unusually strong potential for local adaptation in a marine fish. As part of the California Conservation Genomics Project (CCGP), we sequenced the genome of the...

CaliPopGen is a database of population genetic data for native and naturalized eukaryotic species in California, USA. It summarizes the published literature (1985–2020) for 5,453 unique populations with genetic data from more than 187,394 individuals and 448 species (513 species plus subspecies) across molecular markers including allozymes, RFLPs,...

Keystone species are known to play a critical role in kelp forest health, including the well-known killer whales,sea otter,sea urchin,kelp trophic cascade in the Aleutian Islands, Alaska, USA. In California, a major player in the regulation of sea urchin abundance, and in turn, the health of kelp forests ecosystems, is a large wrasse, the Californi...

The mountains of southern California represent unique, isolated ecosystems that support distinct high-elevation habitats found nowhere else in the area. Analyses of several moisture-dependent species across these sky-islands indicate they exist as locally endemic lineages that are distributed across these fragmented mountains ranges. The Rubber Boa...

The northwestern pond turtle, Actinemys marmorata, and its recently recognized sister species, the southwestern pond turtle, A. pallida, are the sole aquatic testudines occurring over most of western North America, and the only living representatives of the genus Actinemys. Although it historically ranged from Washington state through central Calif...

Dispersal drives invasion dynamics of nonnative species and pathogens. Applying knowledge of dispersal to optimize the management of invasions can mean the difference between a failed and a successful control program and dramatically improve the return on investment of control efforts. A common approach to identifying optimal management solutions f...

The California Conservation Genomics Project (CCGP) is a unique, critically important step forward in the use of comprehensive landscape genetic data to modernize natural resource management at a regional scale. We describe the CCGP, including all aspects of project administration, data collection, current progress, and future challenges. The CCGP...

The past century has witnessed an explosion of anthropogenic activity, resulting in land use and climate changes on a global scale. The study of butterflies provides a unique window into the biological impacts of these changes. In this chapter, we explore several case studies that demonstrate the power of butterflies, both as model organisms in the...

Toxin evolution in animals is one of the most fascinating and complex subjects of scientific inquiry today. Gaining an understanding of toxins poses a multifaceted challenge given the diverse modes of acquisition, evolutionary adaptations, and abiotic components that affect toxin phenotypes. Here, we highlight some of the main genetic and ecologica...

November 2020 marked 2 y since the launch of the Earth BioGenome Project (EBP), which aims to sequence all known eukaryotic species in a 10-y timeframe. Since then, significant progress has been made across all aspects of the EBP roadmap, as outlined in the 2018 article describing the project’s goals, strategies, and challenges (1). The launch phas...

Sea turtles represent an ancient lineage of marine vertebrates that evolved from terrestrial ancestors over 100 MYA, yet the genomic basis of the unique physiological and ecological traits enabling these species to thrive in diverse marine habitats remain largely unknown. Additionally, many populations have declined drastically due to anthropogenic...

Arctostaphylos (Ericaceae) species, commonly known as manzanitas, are an invaluable fire-adapted chaparral clade in the California Floristic Province (CFP), a world biodiversity hotspot on the west coast of North America. This diverse woody genus includes many rare and/or endangered taxa, and the genus plays essential ecological roles in native eco...

This is our 9th edition of an annotated checklist and atlas of all recognized taxa of the world’s modern turtle and tortoise fauna, documenting recent changes and controversies through mid-2021, and including all primary synonyms, updated from eight previous checklists. We provide an updated comprehensive listing of taxonomy and nomenclature, inclu...

As genomic-scale data sets become economically feasible for most organisms, a key question for conservation biology is whether the increased resolution offered by new genomic approaches justifies repeating earlier studies based on traditional markers, rather than investing those same time and monetary resources in less-known species. Genomic studie...

Accurate status assessments of long-lived, widely distributed taxa depend on the availability of long-term monitoring data from multiple populations. However, monitoring populations across large temporal and spatial scales is often beyond the scope of any one researcher or research group. Consequently, wildlife managers may be tasked with utilizing...

Proteins encoded by Antigen Processing Genes (APGs) provide MHC class I (MHC-I) with antigenic peptides. In mammals, polymorphic multigenic MHC-I family is served by monomorphic APGs, whereas in certain non-mammalian species both MHC-I and APGs are polymorphic and coevolve within stable haplotypes. Coevolution was suggested as an ancestral gnathost...

Tetrodotoxin (TTX) is a potent neurotoxin that was first identified in pufferfish but has since been isolated from an array of taxa that host TTX-producing bacteria. However, determining its origin, ecosystem roles, and biomedical applications has challenged researchers for decades. Recognized as a poison and for its lethal effects on humans when i...

Hansson et al . argue that our main finding could provide an overly simplistic metric for maximizing genetic rescue. They agree that translocating the most genetically diverse individuals led to a large increase in translocated tortoise survival, but recommend instead moving individuals that have low genetic load and the greatest representation of...

Hedrick brings up several potential concerns that he feels challenge or limit our main finding. Hedrick does not comment on our empirical results, but rather argues that several factors may confound or invalidate our conclusion. Many of these concerns focus on unknown ecological aspects of the translocated tortoises, but we believe there is no reas...

The western pond turtle (WPT) was formerly considered a single species ( Actinemys or Emys marmorata ) that ranged from southern British Columbia, Canada to Baja California, México. More recently it was divided into a northern and a southern species. WPTs are found primarily in streams that drain into the Pacific Ocean, although scattered populatio...

Significance Population structure and speciation are shaped by a combination of biotic and abiotic factors. The tiger salamander complex has been considered a key group in which life history variation has led to a rapid rate of speciation, driven in large part by the evolution of obligate paedomorphosis—a condition in which adults maintain an aquat...

1. Globally endangered ecosystems, such as ephemeral wetlands, are often critical habitat for multiple interacting imperilled species. To conserve this biodiversity, managers must consider both species-specific resource requirements and mechanisms for endangered species coexistence under variable habitat conditions. 2. We examined communities nati...

As the biodiversity crisis accelerates, the stakes are higher for threatened plants and animals. Rebuilding the health of our planet will require addressing underlying threats at many scales, including habitat loss and climate change. Conservation interventions such as habitat protection, management, restoration, predator control, translocation, ge...

Significance Biodiversity is unevenly distributed across the tree of life. Understanding the factors that led to this unevenness can illuminate how macroevolutionary processes have interacted with changing global environments to shape patterns of biodiversity. By developing a comprehensive phylogeny for extant turtles and analyzing the diversificat...

The phylogeny of California Slender Salamanders (Batrachoseps attenuatus), which inhabits Northern California's Coast Ranges and Sierra Nevada foothills, was previously investigated by Martínez-Solano et al. (2007), who recovered a monophyletic species that contained five geographically cohesive major clades. However, a population isolate of B. att...

Conservation planners use a variety of decision-making tools, many of which require identifying and prioritizing spatial units based on their biodiversity and levels of imperilment. Turtles are highly imperiled, but present schemes for determining global priority areas are focused mostly on broad regional scales. We conduct the first global evaluat...

Hybridization between native and non‐native species is an ongoing global conservation threat. Hybrids that exhibit traits and tolerances that surpass parental values are of particular concern, given their potential to outperform native species. Effective management of hybrid populations requires an understanding of both physiological performance an...

Survival of the most variable As more species become highly threatened because of human activity, there has been an increasing push to understand how best to reintroduce or translocate individuals from wild or captive populations. Suggestions have varied from choosing individuals from the most environmentally similar regions to choosing those that...

Populations of the western spadefoot (Spea hammondii) in southern California occur in one of the most urbanized and fragmented landscapes on the planet and have lost up to 80% of their native habitat. Orange County is one of the last strongholds for this pond-breeding amphibian in the region, and ongoing restoration efforts targeting S. hammondii h...

The North American tiger salamander species complex, including its flagship species the axolotl, has long been a source of biological fascination. The complex exhibits a wide range of variation in developmental life history strategies, including populations and individuals that undergo metamorphosis and those able to forego metamorphosis and retain...

A bstract The North American tiger salamander species complex, including its best-known species, the Mexican axolotl, has long been a source of biological fascination. The complex exhibits a wide range of variation in developmental life history strategies, including populations and individuals that undergo metamorphosis, those able to forego metamo...

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 th...

The western pond turtle (WPT), recently separated into two paripatrically distributed species ( Emys pallida and Emys marmorata ), is experiencing significant reductions in its range and population size. In addition to habitat loss, two potential causes of decline are female-biased road mortality and high juvenile mortality from non-native predator...

Climate change-induced extinctions are estimated to eliminate one in six known species by the end of the century. One major factor that will contribute to these extinctions is extreme climatic events. Here, we show the ecological impacts of recent record warm air temperatures and simultaneous peak drought conditions in California. From 2008–2016, t...

Hybridization between native and non-native species is an ongoing global conservation threat. Hybrids that exhibit traits and tolerances that surpass parental values are of particular concern, given their ability to outcompete the native parent. It is crucial to understand the mechanisms that drive these transgressive hybrid traits to diagnose and...

The barrier effect is a restriction of gene flow between diverged populations by barrier genes. Asymmetric introgression can point to selection, hybrid zone movement, asymmetric reproductive isolation, or a combination of these. Restriction of gene flow and asymmetric introgression over multiple transects point towards influence of intrinsic (genet...

The red-eared slider turtle (Trachemys scripta elegans; RES) is often considered one of the world's most invasive species. Results from laboratory and mesocosm experiments suggest that introduced RES outcompete native turtles for key ecological resources, but such experiments can overestimate the strength of competition. We report on the first fiel...

Landscape genomic signatures indicate reduced gene flow and forest-associated adaptive divergence in an endangered neotropical turtle Running title: Landscape genomics of a neotropical turtle ABSTRACT Human-induced transformations of ecosystems usually result in fragmented populations subject to increased extinction risk. Fragmentation is also ofte...

Genomic data increasingly are used for high resolution population genetic studies including those at the forefront of biological conservation. A key methodological challenge is determining sequence similarity clustering thresholds for RADseq data when no reference genome is available. These thresholds define the maximum permitted divergence among a...

Human‐induced transformations of ecosystems usually result in fragmented populations subject to increased extinction risk. Fragmentation is also often associated with novel environmental heterogeneity, which in combination with restricted gene flow may increase the opportunity for local adaptation. To manage at‐risk populations in these landscapes,...

Newts of the genus Triturus (marbled and crested newts) exhibit substantial variation in the number of trunk vertebrae (NTV) and a higher NTV corresponds to a longer annual aquatic period. Because the Triturus phylogeny has thwarted resolution to date, the evolutionary history of NTV, annual aquatic period, and their potential coevolution has remai...

We present a review and analysis of the conservation status and International Union for Conservation of Nature (IUCN) threat categories of all 360 currently recognized species of extant and recently extinct turtles and tortoises (Order Testudines). Our analysis is based on the 2018 IUCN Red List status of 251 listed species, augmented by provisiona...

Urban biodiversity is an unplanned species assemblage. Although promoting native biodiversity should be the primary goal, the built environment often contains optimal habitat for non-natives. With planning and research, we could use cities as semi-natural assurance colonies for endangered species.

Cryptic species are a challenge for systematics, but their elucidation also may leave critical information gaps about the distribution, conservation status, and behavior of affected species. We use the leopard frogs of the eastern U.S. as a case study of this issue. We refined the known range of the recently described Rana kauffeldi, the Atlantic C...

Cryptic species are a challenge for systematics, but their elucidation also may leave critical information gaps about the distribution, conservation status, and behavior of affected species. We use the leopard frogs of the eastern U.S. as a case study of this issue. We refined the known range of the recently described Rana kauffeldi, the Atlantic C...

Reticulum characterization with ImageJ. (DOCX)

Exemplar photographs of characters used in analysis of leopard frog morphology and patterning. (DOCX)