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The Amphibian Research and Monitoring Initiative (ARMI): 5-year report

The Amphibian Research and Monitoring Initiative (ARMI) is an innovative, multidisciplinary program that began in 2000 in response to a congressional directive for the Department of the Interior to address the issue of amphibian declines in the United States. ARMI’s formulation was cross-disciplinary, integrating U.S. Geological Survey scientists from Biology, Water, and Geography to develop a course of action (Corn and others, 2005a). The result has been an effective program with diverse, yet complementary, expertise. ARMI’s approach to research and monitoring is multiscale. Detailed investigations focus on a few species at selected local sites throughout the country; monitoring addresses a larger number of species over broader areas (typically, National Parks and National Wildlife Refuges); and inventories to document species occurrence are conducted more extensively across the landscape. Where monitoring is conducted, the emphasis is on an ability to draw statistically defensible conclusions about the status of amphibians. To achieve this objective, ARMI has instituted a monitoring response variable that has nationwide applicability. At research sites, ARMI focuses on studying species/environment interactions, determining causes of observed declines, and developing new techniques to sample populations and analyze data. Results from activities at all scales are provided to scientists, land managers, and policymakers, as appropriate. The ARMI program and the scientists involved contribute significantly to understanding amphibian declines at local, regional, national, and international levels. Within National Parks and National Wildlife Refuges, findings help land managers make decisions applicable to amphibian conservation. For example, the National Park Service (NPS) selected amphibians as a vital sign for several of their monitoring networks, and ARMI scientists provide information and assistance in developing monitoring methods for this NPS effort. At the national level, ARMI has had major exposure at a variety of meetings, including a dedicated symposium at the 2004 joint meetings of the Herpetologists’ League, the American Society of Ichthyologists and Herpetologists, and the Society for the Study of Amphibians and Reptiles. Several principal investigators have brought international exposure to ARMI through venues such as the World Congress of Herpetology in South Africa in 2005 (invited presentation by Dr. Gary Fellers), the Global Amphibian Summit, sponsored by the International Union for Conservation of Nature (IUCN) and Wildlife Conservation International, in Washington, D.C., 2005 (invited participation by Dr. P.S. Corn), and a special issue of the international herpetological journal Alytes focused on ARMI in 2004 (edited by Dr. C.K. Dodd, Jr.). ARMI research and monitoring efforts have addressed at least 7 of the 21 Threatened and Endangered Species listed by the U.S. Fish and Wildlife Service (California red-legged frog [Rana draytonii], Chiricahua leopard frog [R. chiricahuensis], arroyo toad [Bufo californicus], dusky gopher frog [Rana sevosa], mountain yellow-legged frog [R. muscosa], flatwoods salamander [Ambystoma cingulatum], and the golden coqui [Eleutherodactylus jasperi]), and 9 additional species of concern recognized by the IUCN. ARMI investigations have addressed time-sensitive research, such as emerging infectious diseases and effects on amphibians related to natural disasters like wildfire, hurricanes, and debris flows, and the effects of more constant, environmental change, like urban expansion, road development, and the use of pesticides. Over the last 5 years, ARMI has partnered with an extensive list of government, academic, and private entities. These partnerships have been fruitful and have assisted ARMI in developing new field protocols and analytic tools, in using and refining emerging technologies to improve accuracy and efficiency of data handling, in conducting amphibian disease, malformation, and environmental effects research, and in implementing a network of monitoring and research sites. Accomplishments from these endeavors include more than 40 publications on amphibian status and trends, nearly 100 publications on amphibian ecology and causes of declines, and over 30 methodological publications. Several databases have emerged as a result of ARMI and its partnerships; one, a digital atlas of ranges for all U.S. amphibian species, was used by the IUCN to display amphibian distribution maps in the Global Amphibian Assessment Project. Given the scope of ARMI and the panoply of projects, findings have had implications for policy. Investigations that demonstrate amphibian declines or illuminate causes of declines provide valuable information about habitat management, environmental effects, mechanisms for the spread of disease, and human/amphibian interfaces. This information has been made available to land managers, scientists, educators, Congress and other policymakers, and the public. The support afforded ARMI by Congress has been influential in the program’s development and success. The value of ARMI’s efforts will continue to increase as we are able to extend our studies spatially and temporally to answer critical questions with more confidence. We are using ARMI’s resources efficiently and continuing to develop innovative mechanisms for leveraging resources for maximum effectiveness during challenging financial times. This report is a 5-year retrospective of the structure, methodology, progress, and contributions to the broader scientific community that have resulted from this national USGS program. We evaluate ARMI’s success to date, with regard to the challenges faced by the program and the strengths that have emerged. We chart objectives for the next 5 years that build on current accomplishments, highlight areas meriting further research, and direct efforts to overcome existing weaknesses.

Scientific Investigations Report

Egg retention in wild-caught Python bivittatus in the Greater Everglades Ecosystem, Florida, USA

Retention of eggs in oviducts beyond the normal oviposition period is a common problem for captive reptiles, but the occurrence of egg retention in wild populations is largely unknown. The Burmese python ( Python [molurus] bivittatus ; Kühl 1820) is an oviparous snake native to south-eastern Asia that is now established in southern Florida. From 2011–2019, invasive Burmese pythons were opportunistically removed from Everglades National Park and Big Cypress National Preserve, humanely euthanised, and necropsied to determine reproductive condition. A total of 258 females of reproductive size were found to exhibit various stages of oviposition which generally aligned with purported annual reproductive timing. However, we encountered five pythons during the post-ovulatory period (Aug–Feb) showing signs of recent oviposition with retained eggs. Most of these cases comprised a small number of retained eggs, likely representing some portion of the total clutch. Because this condition is nearly absent in wild animal literature, our observations suggest retained eggs in wild snakes may be more common than previously assumed, possibly slowing or otherwise impacting population growth. However, we recognise that for an invasive species like the Burmese python in Florida, the egg retention rate may be higher in the non-native range compared to the native range due to maladaptation to novel habitats or environmental conditions. Additional research is needed to determine the exact causes of egg retention and investigate the implications for population dynamics on this and other snake species.

Florida

The effects of geography, habitat, and humans on the ecology and demography of the Gopher tortoise ( Gopherus polyphemus ) in the southern Lake Wales Ridge region of Florida

A 35-year (1967–2002) demographic study was conducted on the gopher tortoise (Gopherus polyphemus) from two different habitats on Archbold Biological Station located on the southern end of the Lake Wales Ridge in south-central Florida. We found geographic, habitat, and human-mediated effects on several aspects of its biology. Our findings underscore the necessity of long-term demographic data to more accurately answer ecological questions concerning long-lived species, such as how the gopher tortoise detectably might be affected by habitat quality and human activities.

Florida

Micro-geographic variation in burrow use of Agassiz’s desert tortoises in the Sonoran Desert of California

Little has been published regarding the burrowing habits of Agassiz’s desert tortoises ( Gopherus agassizii ) in the Sonoran Desert of California. We monitored the interactions of tortoises with their burrows, and other tortoises, via radio-telemetry at two nearby sites between the Cottonwood and Orocopia Mountains, from 2015-2018. We examined how annual cycles of drought and non-drought years, behaviourally affected how tortoises use their burrows (i.e., burrow fidelity, cohabitation, and location), including the timing of the tortoise brumation period. Burrow locations were strongly dependent on local geology and topography, with a tendency to orientate in conformance with the general aspect of the landscape. The timing of brumation was similar to records for G. agassizii throughout their range (with a few exceptions). There was no difference in the estimated number of burrows used per 30 days between the active seasons (2017 and 2018) at the Orocopia site, despite the occurrence of drought in 2018.

California

Assessing effects of pesticides on amphibians and reptiles: status and needs

Growing concern about the decline of certain amphibian populations and for conservation of amphibians and reptiles has led to renewed awareness of problems from pesticides. Testing amphibians and reptiles as a requirement for chemical registration has been proposed but is difficult because of the phylogenetic diversity of these groups. Information from the literature and research may determine whether amphibians and reptiles are adequately protected by current tests for mammals, birds, and fish. Existing information indicates that amphibians are unpredictably more resistant to certain cholinesterase inhibitors, and more sensitive to two chemicals used in fishery applications than could have been predicted. A single study on one species of lizard suggests that reptiles may be close in sensitivity to mammals and birds. Research on effects of pesticides on amphibians and reptiles should compare responses to currently tested groups and should seek to delineate those taxa and chemicals for which cross-group prediction is not possible. New tests for amphibians and reptiles should rely to the greatest extent possible on existing data bases, and should be designed for maximum economy and minimum harm to test animals. A strategy for developing the needed information is proposed. Good field testing and surveillance of chemicals in use may compensate for failures of predictive evaluations and may ultimately lead to improved tests.

Herpetological Journal

Clutch and egg allometry of the turtle Mauremys leprosa (Chelonia: Geoemydidae) from a polluted peri-urban river in west-central Morocco

We examined the relationships of clutch size (CS) and egg size to female body size (straight-line carapace length, CL) in a population of the turtle Mauremys leprosa from a polluted segment of oued (river) Tensift in arid west-central Morocco. Twenty-eight adult females were collected in May–July, 2009 and all were gravid. Each was weighed, measured, humanely euthanized and then dissected. Oviductal shelled eggs were removed, weighed (egg mass, EM) and measured for length (EL) and width (EW). Clutch mass (CM) was the sum of EM for a clutch. Pelvic aperture width (PAW) was measured at the widest point between the ilia bones through which eggs must pass at oviposition. The smallest gravid female had a CL of 124.0 mm. Mean CS was relatively large (9.7±2.0 eggs, range: 3–13) and may reflect high productivity associated with polluted (eutrophic) waters. Regression analyses were conducted using log-transformed data. CM increased isometrically with maternal body size. CS, EW and EM were all significantly hypoallometric in their relationship with CL. EL did not change significantly with increases in CL. EW increased at a hypoallometric rate with increasing CL but was unconstrained by PAW since the widest egg was smaller than the narrowest PAW measurement when excluding the three smallest females. Smaller females may have EW constrained by PAW. As females increase in size they increase both clutch size and egg width in contradiction to predictions of optimal egg size theory.

Tensift River

Comment on Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): request for confirmation of availability of the generic name and for the nomenclatural validation of the journal in which it was published (Case 3601; BZN 70:234–237; 71:30–38; 133-135,181-182 ,252-253)

In Case 3601 Raymond Hoser has asked the Commission to validate for the purposes of nomenclature the name Spracklandus Hoser, 2009, and ‘the journal in which it was published,’ issue 7 of the Australasian Journal of Herpetology (AJH). We note that the entire run of AJH has been written, edited, and published solely by Hoser. Although his requests to the Commission were presented as narrow and, in his words, ‘routine matters,’ we are convinced that they represent an important tipping-point with broad implications of major concern for zoological taxonomy and nomenclature as a whole and, by extension, the greater scientific community. Since Hoser’s actions and works have failed to follow scientific best practices (e.g. Turtle Taxonomy Working Group, 2007, 2014; Kaiser et al., 2013; Kaiser, 2014) and both the Commission’s general Recommendations and Code of Ethics in Appendix A, the global herpetological community has widely rejected his taxonomic decisions and resultant nomenclature. This has unfortunately caused a confusing dual nomenclature to develop in the herpetological community, with most boycotting or ignoring Hoser’s 700+ new names coined in the AJH, while he and a few personal followers actively promote their usage. We believe that suppression of the name Spracklandus , and all issues of AJH, is the only effective way to bring this contentious and confusing issue to resolution. The plenary power available under Article 81.1 of the Code exist specifically to allow the Commission to make rulings in individual cases that disturb stability and cause confusion, whether the works are Code-compliant or not. We maintain that it is in the interest of nomenclatural stability, not only for herpetology, but for all of zoological taxonomy, that the plenary power be invoked to declare the works in AJH unavailable, regardless of any narrow interpretation of their technical Code-compliance. We present our arguments for rejection of the validity of AJH in the following commentary. In view of the wide-reaching implications of this case for all of zoology, and reflecting the deep and broad-based community concern over these issues, our contributing authors include 70 global scientific leaders and accomplished amateurs from a wide variety of zoological disciplines.

Bulletin of Zoological Nomenclature

Habitat use and movement of the endangered Arroyo Toad ( Anaxyrus californicus ) in coastal southern California

Information on the habitat use and movement patterns of Arroyo Toads ( Anaxyrus californicus ) is limited. The temporal and spatial characteristics of terrestrial habitat use, especially as it relates to upland use in coastal areas of the species' range, are poorly understood. We present analyses of radiotelemetry data from 40 individual adult toads tracked at a single site in coastal southern California from March through November of 2004. We quantify adult Arroyo Toad habitat use and movements and interpret results in the context of their life history. We show concentrated activity by both male and female toads along stream terraces during and after breeding, and, although our fall sample size is low, the continued presence of adult toads in the floodplain through the late fall. Adult toads used open sandy flats with sparse vegetation. Home-range size and movement frequency varied as a function of body mass. Observed spatial patterns of movement and habitat use both during and outside of the breeding period as well as available climatological data suggest that overwintering of toads in floodplain habitats of near-coastal areas of southern California may be more common than previously considered. If adult toads are not migrating out of the floodplain at the close of the breeding season but instead overwinter on stream terraces in near-coastal areas, then current management practices that assume toad absence from floodplain habitats may be leaving adult toads over-wintering on stream terraces vulnerable to human disturbance during a time of year when Arroyo Toad mortality is potentially highest.

California

Using spatial capture–recapture to elucidate population processes and space-use in herpetological studies

The cryptic behavior and ecology of herpetofauna make estimating the impacts of environmental change on demography difficult; yet, the ability to measure demographic relationships is essential for elucidating mechanisms leading to the population declines reported for herpetofauna worldwide. Recently developed spatial capture–recapture (SCR) methods are well suited to standard herpetofauna monitoring approaches. Individually identifying animals and their locations allows accurate estimates of population densities and survival. Spatial capture–recapture methods also allow estimation of parameters describing space-use and movement, which generally are expensive or difficult to obtain using other methods. In this paper, we discuss the basic components of SCR models, the available software for conducting analyses, and the experimental designs based on common herpetological survey methods. We then apply SCR models to Red-backed Salamander ( Plethodon cinereus ), to determine differences in density, survival, dispersal, and space-use between adult male and female salamanders. By highlighting the capabilities of SCR, and its advantages compared to traditional methods, we hope to give herpetologists the resource they need to apply SCR in their own systems.

Journal of Herpetology

General herpetological collecting is size-biased for five Pacific lizards

Accurate estimation of a species' size distribution is a key component of characterizing its ecology, evolution, physiology, and demography. We compared the body size distributions of five Pacific lizards ( Carlia ailanpalai, Emoia caeruleocauda, Gehyra mutilata, Hemidactylus frenatus , and Lepidodactylus lugubris ) from general herpetological collecting (including visual surveys and glue boards) with those from complete censuses obtained by total removal. All species exhibited the same pattern: general herpetological collecting undersampled juveniles and oversampled mid-sized adults. The bias was greatest for the smallest juveniles and was not statistically evident for newly maturing and very large adults. All of the true size distributions of these continuously breeding species were skewed heavily toward juveniles, more so than the detections obtained from general collecting. A strongly skewed size distribution is not well characterized by the mean or maximum, though those are the statistics routinely reported for species' sizes. We found body mass to be distributed more symmetrically than was snout–vent length, providing an additional rationale for collecting and reporting that size measure.

Mariana Islands

Making great leaps forward: Accounting for detectability in herpetological field studies

Detecting individuals of amphibian and reptile species can be a daunting task. Detection can be hindered by various factors such as cryptic behavior, color patterns, or observer experience. These factors complicate the estimation of state variables of interest (e.g., abundance, occupancy, species richness) as well as the vital rates that induce changes in these state variables (e.g., survival probabilities for abundance; extinction probabilities for occupancy). Although ad hoc methods (e.g., counts uncorrected for detection, return rates) typically perform poorly in the face of no detection, they continue to be used extensively in various fields, including herpetology. However, formal approaches that estimate and account for the probability of detection, such as capture-mark-recapture (CMR) methods and distance sampling, are available. In this paper, we present classical approaches and recent advances in methods accounting for detectability that are particularly pertinent for herpetological data sets. Through examples, we illustrate the use of several methods, discuss their performance compared to that of ad hoc methods, and we suggest available software to perform these analyses. The methods we discuss control for imperfect detection and reduce bias in estimates of demographic parameters such as population size, survival, or, at other levels of biological organization, species occurrence. Among these methods, recently developed approaches that no longer require marked or resighted individuals should be particularly of interest to field herpetologists. We hope that our effort will encourage practitioners to implement some of the estimation methods presented herein instead of relying on ad hoc methods that make more limiting assumptions.

Journal of Herpetology

Time-to-detection occupancy modeling: An efficient method for analyzing the occurrence of amphibians and reptiles

Occupancy models provide a reliable method of estimating species distributions while accounting for imperfect detectability. The cost of accounting for false absences is that detection and nondetection surveys typically require repeated visits to a site or multiple-observer techniques. More efficient methods of collecting data to estimate detection probabilities would allow additional sites to be surveyed for the same amount of effort, which would support more precise estimation of covariate effects to improve inference about underlying ecological processes. Time-to-detection surveys allow the estimation of detection probability based on a single site visit by one observer, and therefore might be an efficient technique for herpetological occupancy studies. We evaluated the use of time-to-detection surveys to estimate the occupancy of pond-breeding amphibians at Point Reyes National Seashore, California, USA, including variables that affected detection rates and the probability of occurrence. We found that detection times were short enough, and occupancy was high enough, to estimate reliably the probability of occurrence of three pond-breeding amphibians at Point Reyes National Seashore, and that survey and site conditions had species-specific effects on detection rates. In particular, pond characteristics affected detection times of all commonly detected species. Probability of occurrence of Sierran Treefrogs ( Hyliola sierra ) and Rough-Skinned Newts ( Taricha granulosa ) was negatively related to the detection of fish and pond area. Time-to-detection surveys can provide an efficient method for estimating detection probabilities and accounting for false absences in occupancy studies of reptiles and amphibians.

California

Population structure of the African Clawed Frog (Xenopus laevis) in maize-growing areas with atrazine application versus non-maize-growing areas in South Africa

The herbicide atrazine has been suggested to cause gonadal deformities in frogs and could possibly impact on reproduction. Since the early 1960s, atrazine has been used in large amounts in maize production areas of South Africa. These areas overlap with populations of the African Clawed Frog (Xenopus laevis) that has a wide distribution in southern Africa and is found in most water-bodies including those where atrazine residues are detected. The aim of this study was to compare various attributes of individual- and population-level responses of X. laevis from maize-growing and non-maize-growing areas. Xenopus laevis were studied in three reference and five maize-growing sites. Sex ratio, snout-vent length, body-mass and age profiles were found to be similar for populations in maize-growing and non-maize-growing areas. Our mark-recapture data indicated that all sites had robust populations. There were no significant relationships between exposure to atrazine and any of the parameters investigated in populations of X. laevis.

African Journal of Herpetology

Assessment of laryngeal muscle and testicular cell types in Xenopus laevis (Anura Pipidae) inhabiting maize and non-maize growing areas of South Africa

We tested the hypothesis that adult African clawed frogs (Xenopus laevis) inhabiting water bodies in maize-growing areas (MGA) of South Africa would exhibit differences in testicular structure compared to frogs from water bodies in non-maize-growing areas (NMGA) in the same locale. Adults of both sexes were collected during the autumn of 2002 in South Africa, and stereological analytical techniques were used to quantify the distribution of testicular cell types. In addition, total laryngeal mass was used as a gauge of secondary sex differences in animals from MGA and NMGA study sites. Evaluation of the total laryngeal mass revealed that there were no statistically significant differences between X. laevis of the same sex from the NMGA and MGA sites. Mean percent fractional-volume values for seminiferous tubule distribution of testicular cell types of mature X. laevis, ranged from 3-4% for spermatogonia, 26-28% for spermatocytes, 54-57% for spermatozoa, and 14-15% for other cells types. The mean percent volume for blood vessels ranged from 0.3-0.4%. These values did not differ significantly between frogs from NMGA and MGA areas. Collectively, these data demonstrated no differences in gonadal and laryngeal development in X. laevis collected in South Africa from MGA and NMGA areas and that there is little evidence for an effect of agricultural chemicals used in maize production functioning as endocrine disrupters in this species. Screening of X. laevis testes revealed a small incidence of Stage 1 testicular oocytes in adult male frogs collected from the NMGA (3%) and MGA (2%).

African Journal of Herpetology

Population dynamics of Eleutherodactylus coqui in cordillera forest reserves of Puerto Rico

Various aspects of population structure and dynamics of Eleutherodactylus coqui in two forest reserves (Maricao and Guilarte) of the central mountain range of Puerto Rico were determined between July 1997 and June 1998. Adult density ranged from 8–25 animals/100 m 2 for the wet season and 3–19 animals/100 m 2 for the dry season. Abundance of froglets and juveniles was also greater in the wet season compared to the dry season. By contrast, egg mass counts were greater in the dry season compared to the wet season. Adult body size increased from wet to dry season as population density declined and was found to differ significantly between forests. Adult E. coqui trapped at similar elevations were larger in Guilarte compared to those in Maricao. Average body size for Maricao and Guilarte populations were smaller than those reported for populations in eastern Puerto Rico. Comparing average snout–vent length among frogs from the Luquillo Mountains, Guilarte Forest, and Maricao Forest, a gradient of decreasing body size was observed from east to west across the island.

Puerto Rico