Empirical evidence for effects of invasive American Bullfrogs on occurrence of native amphibians and emerging pathogens.

Invasive species and emerging infectious diseases are two of the greatest threats to biodiversity. American Bullfrogs (Rana [Lithobates] catesbeiana), which have been introduced to many parts of the world, are often linked with declines in native amphibians via predation and the spread of emerging pathogens such as amphibian chytrid fungus (Batrachochytrium dendrobatidis [Bd]) and ranaviruses. Although many studies have investigated the potential role of bullfrogs in the decline of native amphibians, analyses that account for shared habitat affinities and imperfect detection have found limited support for clear effects. Similarly, the role of bullfrogs in shaping the patch-level distribution of pathogens is unclear. We used eDNA methods to sample 233 sites in the southwestern USA and Sonora, Mexico (2016-2018) to estimate how the presence of bullfrogs affects the occurrence of four native amphibians, Bd, and ranaviruses. Based on two-species, dominant-subordinate occupancy models fitted in a Bayesian context, federally threatened Chiricahua Leopard Frogs (Rana chiricahuensis) and Western Tiger Salamanders (Ambystoma mavortium) were eight times (32% vs. 4%) and two times (36% vs. 18%), respectively, less likely to occur at sites where bullfrogs occurred. Evidence for the negative effects of bullfrogs on Lowland Leopard Frogs (Rana yavapaiensis) and Northern Leopard Frogs (Rana pipiens) was less clear, possibly because of smaller numbers of sites where these native species still occurred and because bullfrogs often occur at lower densities in streams, the primary habitat for Lowland Leopard Frogs. At the community level, Bd was most likely to occur where bullfrogs co-occurred with native amphibians, which could increase the risk to native species. Ranaviruses were estimated to occur at 33% of bullfrog-only sites, 10% of sites where bullfrogs and native amphibians co-occurred, and only 3% of sites where only native amphibians occurred. Of the 85 sites where we did not detect any of the five target amphibian species, we also did not detect Bd or ranaviruses; this suggests other hosts do not drive the distribution of these pathogens in our study area. Our results provide landscape-scale evidence that bullfrogs reduce the occurrence of native amphibians and increase the occurrence of pathogens, information that can clarify risks and aid the prioritization of conservation actions.

Factors related to species richness, endemism, and conservation status of the herpetofauna (Amphibia and Reptilia) of Mexican states.

Mexico is a megadiverse country with high endemicity in its herpetofauna. We examine how species richness, proportion of state and country endemic species, and proportion of species in a category of conservation concern using listings in the International Union for Conservation of Nature (IUCN) Red List and the Secretaría del Medio Ambiente y Recursos Naturales (SEMARNAT) in 27 of 32 Mexican states are related to environmental and human demographic and socioeconomic variables. Amphibian and reptile species richness were positively related to latitude range and number of physiographic regions and negatively related to latitude. The proportion of state endemic amphibian species in a state was negatively related to latitude whereas no variables influenced the proportion in reptiles. The proportion of country endemics in a state was positively related to human population density and the number of physiographic regions and negatively related to per capita gross domestic product (GDP) and latitude range for amphibians; it was positively related to human population density and elevation range and negatively related to latitude range for reptiles. The proportion of amphibian species in an IUCN category of concern in a state was positively related to human population density and negatively related to latitude; for reptiles, it was negatively related to human population density. The proportion of SEMARNAT-listed species in a state was positively related to human population density for both amphibians and reptiles and negatively related to latitude range for amphibians. Our analyses found that larger macroecological patterns (e.g., latitudinal species gradient, heterogeneity-richness relationships) and human population density play important roles in determining the richness and conservation status of Mexican amphibians and reptiles.

A conservation checklist of the amphibians and reptiles of the State of Mexico, Mexico with comparisons with adjoining states.

The State of Mexico has a unique combination of geographic characteristics and topography that promotes a high biodiversity. Unfortunately, continued human population growth of the metropolitan areas of Mexico City and Toluca have degraded the environment of the State of Mexico, which threatened its wildlife. An updated checklist of the amphibians and reptiles of the State of Mexico is provided and their conservation status summarized. The State of Mexico has 49 species of amphibians and 101 species of reptiles. The majority of the amphibians (73.5%) and reptiles (70.3%) found in the State of Mexico are endemic to Mexico. Of the amphibian and reptile species in the State of Mexico, 20.1% are IUCN listed (i.e., Vulnerable, Near Threatened, or Endangered), 18.4% are placed in a protected category by SEMARNAT (excluding NL and Pr, this last category is equivalent to the LC category of IUCN), and 34.9% are categorized as high risk by the EVS. The importance of forested habitats for the protected amphibians and reptiles in the State of Mexico suggest that management of these habitats to maintain or expand them needs to be considered.

A conservation checklist of the amphibians and reptiles of Mexico City, with comparisons with adjoining states.

Mexico City houses one of the most populous urban areas of the world, and the modification of its natural habitat likely influences the biological diversity found there. In particular, amphibians and reptiles are likely affected by these modifications. Herein, we present an updated list of the species of amphibians and reptiles that inhabit Mexico City. Mexico City harbors 65 species of amphibians and reptiles, which represent 21 families and 33 genera. These include 18 species of amphibians (nine anurans and nine salamanders) and 47 species of reptiles (14 lizards, 30 snakes [one introduced], and three turtles [one introduced]). Forty-eight of the amphibian and reptile species in Mexico City are endemic to Mexico, with two endemic to Mexico City. The most diverse region of Mexico City is the Forests and Ravines region, which is home to 43 species. Eleven species of amphibians and reptiles in Mexico City are IUCN listed, 16 are placed in a protected category by SEMARNAT (Secretaria del Medio Ambiente y Recursos Naturales), and 27 species are categorized as high risk by the EVS (Environmental Viability Score). Mexico City shares almost 94% of its species with the State of Mexico.

Batrachochytrium salamandrivorans (Bsal) not detected in an intensive survey of wild North American amphibians.

The salamander chytrid fungus (Batrachochytrium salamandrivorans [Bsal]) is causing massive mortality of salamanders in Europe. The potential for spread via international trade into North America and the high diversity of salamanders has catalyzed concern about Bsal in the U.S. Surveillance programs for invading pathogens must initially meet challenges that include low rates of occurrence on the landscape, low prevalence at a site, and imperfect detection of the diagnostic tests. We implemented a large-scale survey to determine if Bsal was present in North America designed to target taxa and localities where Bsal was determined highest risk to be present based on species susceptibility and geography. Our analysis included a Bayesian model to estimate the probability of occurrence of Bsal given our prior knowledge of the occurrence and prevalence of the pathogen. We failed to detect Bsal in any of 11,189 samples from 594 sites in 223 counties within 35 U.S. states and one site in Mexico. Our modeling indicates that Bsal is highly unlikely to occur within wild amphibians in the U.S. and suggests that the best proactive response is to continue mitigation efforts against the introduction and establishment of the disease and to develop plans to reduce impacts should Bsal establish.

A conservation checklist of the herpetofauna of Morelos, with comparisons with adjoining states.

Despite being one of the smallest states in Mexico, the high diversity of habitats in Morelos has led to the development of a rich biota made up of a mixture of species typical of the Neovolcanic Axis and the Sierra Madre del Sur. However, recent expansion of cities in Morelos is likely to have consequences for the state's herpetofauna. Here a checklist of the amphibians and reptiles of Morelos is provided with a summary of their conservation status and overlap with its neighboring states. Morelos is home to 139 species of amphibians and reptiles representing 32 families and 75 genera. Twenty-six of the 38 species of amphibians and 70 of the 101 species of reptiles that inhabit Morelos are endemic to Mexico. Fourteen species of amphibians and reptiles from Morelos are IUCN listed (i.e., Vulnerable, Near Threatened, or Endangered), 22 are placed in a protected category by SEMARNAT, and 41 are categorized as high risk by the EVS. The Tropical Deciduous Forest vegetation type hosts the greatest number of amphibian and reptile species in Morelos (84 species). Morelos shares the largest proportion of its herpetofauna with the State of Mexico (79.3%), Puebla (77.0%), and Guerrero (74.8%).

The amphibians and reptiles of Colima, Mexico, with a summary of their conservation status.

Colima is the fourth smallest Mexican state, covering only 0.3% of the surface area of Mexico, but due to the remarkable diversity of physiographic and environmental conditions present in Colima it contains a high biological diversity. We generated an up-to-date herpetofaunal checklist for Colima, with a summary of the conservation status of Colima's amphibians and reptiles. Our checklist contains a total of 153 species of amphibians and reptiles (three introduced). Thirty-nine are amphibians and 114 are reptiles. More than half of Colima's herpetofauna are Mexican endemics (66.7% of amphibians, 67.5% of reptiles). Less than 25% of the amphibian and reptile species in Colima are in protected categories according to the IUCN Red List and SEMARNAT. The reptiles in the Marine and Revillagigedo Archipelago regions are the most threatened taxa of the Colima herpetofauna. Colima shares > 80% of its herpetofauna with its neighboring states, Jalisco and Michoacán.

A checklist of the amphibians and reptiles of Sinaloa, Mexico with a conservation status summary and comparisons with neighboring states.

Sinaloa possesses a rich biota with unique characteristics due to its proximity to the northern deserts, the tropical lowlands of the south, and the temperate environments of the western slopes of the Sierra Madre Occidental in Mexico. However, threats to its environment makes understanding the biological diversity of Sinaloa crucial. A checklist of the amphibians and reptiles has been generated, and the conservation status of Sinaloa's herpetofauna summarized with the aim of understanding the potential conservation or management needs. Sinaloa has 159 species of amphibians and reptiles, including 39 species of amphibians and 120 species of reptiles. The herpetofauna of Sinaloa has relatively few species of conservation concern at a global and national scale (IUCN and SEMARNAT lists), but Environmental Vulnerability Scores suggest that there might be greater conservation concerns for the Sinaloa herpetofauna. Families of particular conservation concern include Craugastoridae, Eleutherodactylidae, Ambystomatidae, Crocodylidae, Dactyloidae, Eublepharidae, Helodermatidae, Iguanidae, Phrynosomatidae, Phyllodactylidae, Colubridae, Natricidae, Viperidae, Cheloniidae, and Dermochelyidae.

Environmental heterogeneity and not vicariant biogeographic barriers generate community-wide population structure in desert-adapted snakes.

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.

A conservation checklist of the amphibians and reptiles of Sonora, Mexico, with updated species lists.

Sonora has a rich natural diversity, including reptiles and amphibians. Sonora's location on the United States-Mexico border creates some unique conservation challenges for its wildlife. We compiled a list of the amphibian and reptile species currently known for Sonora, summarized the conservation status of these species, and compared our list of species with known species lists for adjacent states. The herpetofauna of Sonora comprises 200 species of amphibians and reptiles (38 amphibians and 162 reptiles). Overall, Sonora shares the most species with Chihuahua, Sinaloa, and Arizona. Approximately 11% of the amphibian and reptile species are IUCN listed, but 35.5% are placed in a protected category by SEMARNAT, and 32.6% are categorized as high risk by the Environmental Vulnerability Score.

Amphibians and reptiles of the state of San Luis Potosí, Mexico, with comparisons with adjoining states.

A summary of the species of amphibians and reptiles of the state has been compiled, including their geographic distributions, habitats, and conservation statuses. The herpetofauna of San Luis Potosí consists of 41 species of amphibians and 141 species of reptiles. San Luis Potosí shares the highest number of species with Hidalgo and Tamaulipas, and the least number of species with Nuevo León. In San Luis Potosí, there are several taxa of particular conservation concern including salamanders, emydid and trionychid turtles, anguid and xenosaurid lizards, and natricid and colubrid snakes.

Amphibians and reptiles of the state of Durango, Mexico, with comparisons with adjoining states.

A summary of the species of amphibians and reptiles of Durango, as well as their geographic distributions, habitat, and conservation status have been compiled. The herpetofauna of Durango consists of 36 species of amphibians and 120 species of reptiles. Durango shares the most species with Chihuahua (74.0%), and shares fewer species with Sinaloa (48.0%), Nayarit (48.7%), and Coahuila (48.0%). Arid-semiarid and Sierras habitat types have the most species, with valleys and Quebradas habitat types having fewer species. In Durango, there are several taxa of particular conservation concern including eleutherodactylid frogs, eublepharid, iguanid, phrynosomatid, and xantusid lizards, boid, colubrid, and natricid snakes, and emydid and testudinid turtles.

Amphibians and reptiles of the state of Chihuahua, Mexico, with comparisons with adjoining states.

Chihuahua is Mexico's largest state, and its physiographic complexity affects the distribution of its herpetofauna. We list amphibians and reptiles for the state of Chihuahua, with their conservation status. We also compare this list to those of six adjoining states in the United States and Mexico (New Mexico, Texas, Coahuila, Durango, Sinaloa, and Sonora). A total of 175 species of amphibians and reptiles is found in Chihuahua. Thirty-eight are amphibians, and 137 reptiles. Chihuahuan amphibians and reptiles represent just over 37% of such species from Chihuahua and neighboring states. Chihuahua shares the highest proportion of its herpetofauna with Sonora and Durango. Most of the herpetofauna of Chihuahua falls in IUCNs least concern category and is not listed by SEMARNAT. However, turtles in Chihuahua are a group of particular conservation concern.

Lineage diversification of fringe-toed lizards (Phrynosomatidae: Uma notata complex) in the Colorado Desert: Delimiting species in the presence of gene flow.

Multi-locus nuclear DNA data were used to delimit species of fringe-toed lizards of the Uma notata complex, which are specialized for living in wind-blown sand habitats in the deserts of southwestern North America, and to infer whether Quaternary glacial cycles or Tertiary geological events were important in shaping the historical biogeography of this group. We analyzed ten nuclear loci collected using Sanger sequencing and genome-wide sequence/single-nucleotide polymorphism (SNP) data collected using restriction-associated DNA (RAD) sequencing. A combination of species discovery methods (concatenated phylogenies, parametric and non-parametric clustering algorithms) and species validation approaches (coalescent-based species tree/isolation-with-migration models) were used to delimit species, infer phylogenetic relationships, and to estimate effective population sizes, migration rates, and speciation times. Uma notata, U. inornata, U. cowlesi, and an undescribed species from Mohawk Dunes, Arizona (U. sp.) were supported as distinct in the concatenated analyses and by clustering algorithms, and all operational taxonomic units were decisively supported as distinct species by ranking hierarchical nested speciation models with Bayes factors based on coalescent-based species tree methods. However, significant unidirectional gene flow (2NM>1) from U. cowlesi and U. notata into U. rufopunctata was detected under the isolation-with-migration model. Therefore, we conservatively delimit four species-level lineages within this complex (U. inornata, U. notata, U. cowlesi, and U. sp.), treating U. rufopunctata as a hybrid population (U. notata×cowlesi). Both concatenated and coalescent-based estimates of speciation times support the hypotheses that speciation within the complex occurred during the late Pleistocene, and that the geological evolution of the Colorado River delta during this period was an important process shaping the observed phylogeographic patterns.

A diversity and conservation inventory of the Herpetofauna of the Cuautlapan Valley, Veracruz, Mexico.

We compiled an inventory of the amphibians and reptiles of the Cuautlapan Valley, Veracruz, Mexico based on field surveys and museum and literature records. We found a total of 78 species: 28 amphibians (6 Salamanders and 22 anurans); and 50 reptiles (three turtles, 18 lizards, and 29 snakes). These taxa represent 26 families (eight amphibian families, 18 reptile families) and 60 genera (19 amphibian genera, 41 reptile genera). Two of these species are not native to the area (Rhinoclemmys pulcherrima and Hemidactylus frenatus). According to the IUCN red list, five species are Critically Endangered, two are Endangered, four are Near Threatened, and four are Vulnerable. In the SEMARNAT (Secretaría del Medio Ambiente y Recursos Naturales) listing, one species is Endangered, eight species are Threatened, and 25 are Subject to Special Protection. Even though the Cuautlapan Valley is represented by a relatively small area it hosts a rich diversity of amphibian and reptile species, many of which are at risk and protected under Mexican law. This valley lies between the growing cities of Orizaba and Cordoba which have contributed to habitat degradation threatening the existence of the wildlife that occurs there.

Amphibians and reptiles of the state of Coahuila, Mexico, with comparison with adjoining states.

We compiled a checklist of the amphibians and reptiles of the state of Coahuila, Mexico. The list comprises 133 species (24 amphibians, 109 reptiles), representing 27 families (9 amphibians, 18 reptiles) and 65 genera (16 amphibians, 49 reptiles). Coahuila has a high richness of lizards in the genus Sceloporus. Coahuila has relatively few state endemics, but has several regional endemics. Overlap in the herpetofauna of Coahuila and bordering states is fairly extensive. Of the 132 species of native amphibians and reptiles, eight are listed as Vulnerable, six as Near Threatened, and six as Endangered in the IUCN Red List. In the SEMARNAT listing, 19 species are Subject to Special Protection, 26 are Threatened, and three are in Danger of Extinction. Coahuila is home to several species of conservation concern, especially lizards and turtles. Coahuila is an important state for the conservation of the native regional fauna.

Amphibians and Reptiles of the state of Nuevo León, Mexico.

We compiled a check list of the herpetofauna of Nuevo León. We documented 132 species (23 amphibians, 109 reptiles), representing 30 families (11 amphibians, 19 reptiles) and 73 genera (17 amphibians, 56 reptiles). Only two species are endemic to Nuevo León. Nuevo León contains a relatively high richness of lizards in the genus Sceloporus. Overlap in the herpetofauna of Nuevo León and states it borders is fairly extensive. Of 130 native species, 102 are considered species of Least Concern in the IUCN red list, four are listed as Vulnerable, five are listed as Near Threatened, and four are listed as Endangered. According to SEMARNAT, 78 species are not of conservation concern, 25 are subject to Special Protection, 27 are Threatened, and none are listed as in Danger of Extinction. Given current threats to the herpetofauna, additional efforts to understand the ecology and status of populations in Nuevo León are needed.

Do gravid female Anolis nebulosus thermoregulate differently than males and non-gravid females?

In lizards, the role of gravid oviparous females in controlling the temperature experienced by developing embryos prior to oviposition has been rarely examined. In particular, relatively little is known about the effect of gravidity on the thermal ecology of female Anolis lizards. Here we examine the thermal ecology of Anolis nebulosus from Nayarit, Mexico, with a particular goal of comparing the thermal ecology of gravid females to that of non-gravid females and males. The thermal efficiency (E) of gravid female A. nebulosus (E=0.782) was higher than in males (E=0.464), and to a lesser extent, non-gravid females (E=0.637), despite no significant differences observed in body, air, operative, or set point temperatures among males, gravid females, and non-gravid females. Gravid females had smaller differences between body temperatures and set point temperatures (db), but did not differ in the difference between operative temperature and set point temperature (de). Gravid females used sun-shade and shaded microhabitats proportionately more than males and non-gravid females, and rarely used sunny microhabitats. Our results suggest that gravid A. nebulosus are using a different and more efficient thermoregulatory strategy than other adults in the population. Such efficient thermoregulation is possibly related to females attempting to provide a thermal environment that is conducive to the development of embryos in eggs prior to oviposition.

A multilocus perspective on the speciation history of a North American aridland toad (Anaxyrus punctatus).

Interpretations of phylogeographic patterns can change when analyses shift from single gene-tree to multilocus coalescent analyses. Using multilocus coalescent approaches, a species tree and divergence times can be estimated from a set of gene trees while accounting for gene-tree stochasticity. We utilized the conceptual strengths of a multilocus coalescent approach coupled with complete range-wide sampling to examine the speciation history of a broadly distributed, North American warm-desert toad, Anaxyrus punctatus. Phylogenetic analyses provided strong support for three major lineages within A. punctatus. Each lineage broadly corresponded to one of three desert regions. Early speciation in A. punctatus appeared linked to late Miocene-Pliocene development of the Baja California peninsula. This event was likely followed by a Pleistocene divergence associated with the separation of the Chihuahuan and Sonoran Deserts. Our multilocus coalescent-based reconstruction provides an informative contrast to previous single gene-tree estimates of the evolutionary history of A. punctatus.

TEMPORAL VARIATION IN THE ABUNDANCE OF Poblana alchichica IN NEAR-SHORE HABITAT OF THE HIGH ELEVATION LAKE, LAGO DE ALCHICHICA, PUEBLA, MEXICO / Variación temporal en la abundancia de Poblana alchichica en un hábitat ribereño del lago de alta elevación, Lago de Alchichica, Puebla, México

The distribution and abundance of organisms within a pond or lake can reflect the result of a variety of factors. We examined the abundance of the endemic fish, Poblana alchichica, in Lago de Alchichica, Puebla, Mexico, as well as how the abundance varied among months. Dissolved oxygen, temperature, and salinity varied among months. The abundance of Poblana alchichica peaked in December and February. For the months when fish were observed, their abundances were positively related to dissolved oxygen concentration and were generally not related to temperature. Our results provide evidence that there is substantial seasonal variation in the abundances of Poblana alchichica and that within months, their distributions are likely driven by dissolved oxygen than either temperature or salinity.

La distribución y abundancia de organismos dentro de un cuerpo de agua puede reflejar el resultado de variedad de factores. Examinamos la abundancia del pez endémico, Poblana alchichica, en el lago de Alchichica, Puebla, México, así como la variación de la abundancia entre los meses del estudio. El oxígeno disuelto, temperatura y salinidad variaron entre los meses. La abundancia de Poblana alchichica fue mayor en diciembre y febrero. Durante los meses en los que se observaron a los peces, su abundancia fue relacionada positivamente con la concentración de oxígeno disuelto, y por lo general no presentaron relación con la temperatura. Nuestros resultados muestran evidencia clara de variación estacional en la abundancia de Poblana alchichica, y cómo su distribución se asocia más probablemente con oxígeno disuelto que con temperatura o salinidad.