Do hormone‐modulating chemicals impact on reproduction and development of wild amphibians?

Globally, amphibians are undergoing a precipitous decline. At the last estimate in 2004, 32% of the approximately 6000 species were threatened with extinction and 43% were experiencing significant declines. These declines have been linked with a wide range of environmental pressures from habitat loss to climate change, disease and pollution. This review evaluates the evidence that endocrine‐disrupting contaminants (EDCs) – pollutants that affect hormone systems – are impacting on wild amphibians and contributing to population declines. The review is limited to anurans (frogs and toads) as data for effects of EDCs on wild urodeles (salamanders, newts) or caecilians (limbless amphibians) are extremely limited. Evidence from laboratory studies has shown that a wide range of chemicals have the ability to alter hormone systems and affect reproductive development and function in anurans, but for the most part only at concentrations exceeding those normally found in natural environments. Exceptions can be found for exposures to the herbicide atrazine and polychlorinated biphenyls in leopard frogs (Rana pipiens) and perchlorate in African clawed frogs (Xenopus laevis). These contaminants induce feminising effects on the male gonads (including ‘intersex’ – oocytes within testes) at concentrations measured in some aquatic environments. The most extensive data for effects of an EDC in wild amphibian populations are for feminising effects of atrazine on male gonad development in regions across the USA. Even where strong evidence has been provided for feminising effects of EDCs, however, the possible impact of these effects on fertility and breeding outcome has not been established, making inference for effects on populations difficult. Laboratory studies have shown that various chemicals, including perchlorate, polychlorinated biphenyls and bromodiphenylethers, also act as endocrine disrupters through interfering with thyroid‐dependent processes that are fundamental for amphibian metamorphosis. Perchlorate has also been shown to induce these effects in wild anuran populations from perchlorate‐contaminated environments. Overall, the published data available suggest that some health effects observed in wild anuran populations, most notably intersex, likely have a chemical aetiology; however they derive only from very few anuran species and for a few pesticides at field sites in the USA. To understand better the impacts of EDCs on wild anuran populations, as well as other amphibian groups, assessment of fertility in exposed animals are required. Development of non‐destructive biomarkers that are indicative of specific EDC‐effect mechanisms are also needed to allow the study of vulnerable populations. This will help to distinguish the effects of EDCs from other environmental and/or genetic influences on development and reproduction.     

Understanding of the impact of chemicals on amphibians: a meta-analytic review

Many studies have assessed the impact of different pollutants on amphibians across a variety of experimental venues (laboratory, mesocosm, and enclosure conditions). Past reviews, using vote-counting methods, have described pollution as one of the major threats faced by amphibians. However, vote-counting methods lack strong statistical power, do not permit one to determine the magnitudes of effects, and do not compare responses among predefined groups. To address these challenges, we conducted a meta-analysis of experimental studies that measured the effects of different chemical pollutants (nitrogenous and phosphorous compounds, pesticides, road deicers, heavy metals, and other wastewater contaminants) at environmentally relevant concentrations on amphibian survival, mass, time to hatching, time to metamorphosis, and frequency of abnormalities. The overall effect size of pollutant exposure was a medium decrease in amphibian survival and mass and a large increase in abnormality frequency. This translates to a 14.3% decrease in survival, a 7.5% decrease in mass, and a 535% increase in abnormality frequency across all studies. In contrast, we found no overall effect of pollutants on time to hatching and time to metamorphosis. We also found that effect sizes differed among experimental venues and among types of pollutants, but we only detected weak differences among amphibian families. These results suggest that variation in sensitivity to contaminants is generally independent of phylogeny. Some publication bias (i.e., selective reporting) was detected, but only for mass and the interaction effect size among stressors. We conclude that the overall impact of pollution on amphibians is moderately to largely negative. This implies that pollutants at environmentally relevant concentrations pose an important threat to amphibians and may play a role in their present global decline.     

Effects of UV-B Radiation on Anti-predator Behavior in Three Species of Amphibians

Ultraviolet radiation has been suggested as a possible contributing cause of amphibian declines around the world. Both laboratory and field studies have demonstrated that exposure to ultraviolet radiation can lead to increased mortality of developing amphibians. Virtually no studies have examined the sub‐lethal effects of ultraviolet on amphibian behavior. In this study, we examine the anti‐predator behavior of three species of amphibians after short‐term exposure to ultraviolet‐B radiation. Toad (Bufo boreas) juveniles that had been exposed to ultraviolet radiation did not respond to chemical extracts from conspecifics and heterospecifics as much as juveniles that had not been exposed. Both newt larvae (Taricha granulosa) that had been exposed to ultraviolet radiation and those that had not been exposed responded to chemical cues from conspecific predators by increasing the amount of time spent in shelter. Frog tadpoles (Rana cascadae) that had been exposed to ultraviolet radiation did not reduce their movement in response to chemical cues from predators as much as tadpoles that had not been exposed. These results indicate that ultraviolet exposure may have important sub‐lethal effects in amphibians that could adversely effect their fitness.     

Burning issues for conservation: A critique of faunal fire research in Southern Africa

Abstract Fire is a key ecological process in several biomes worldwide. Although many conservation agencies have the protection of biodiversity as at least one of their major goals, information on the effects of fire on fauna in these biomes is fragmentary. Here we provide an overview of the published research undertaken to date on the effects of fire on fauna using examples from Southern Africa. We found that few studies have examined the effects of fire on amphibians or reptiles, and work on invertebrates is likewise sparse. The majority of studies that have been published are observational reports, and few experimental studies have been undertaken using an experimental fire regime, or over appropriately long time intervals. Replication was often not reported and where this was done, it was generally inadequate. The majority of the studies failed to report the area over which the studies were undertaken and sampling unit size was often not reported. Despite the importance of fire duration, ignition method, season and time of day of fire, few studies investigated these variables. We conclude that at present the information on the effects of fire on fauna in Southern Africa is fragmentary and, consequently, informed management decisions regarding the consequences of burning policies on the conservation of biodiversity both within and outside protected areas are problematic. We recommend that future studies, both in Southern Africa and elsewhere, be based on a suite of large‐scale and experimental approaches (the latter firmly grounded in the principles of sound experimental design), use non‐classical statistics to explore the effects of large‐scale or unreplicated fires, and where possible include baseline information such as that gathered in fragmentation experiments.     

Amphibians as models for studying environmental change

The use of amphibians as models in ecological research has a rich history. From an early foundation in studies of amphibian natural history sprang generations of scientists who used amphibians as models to address fundamental questions in population and community ecology. More recently, in the wake of an environment that human disturbances rapidly altered, ecologists have adopted amphibians as models for studying applied ecological issues such as habitat loss, pollution, disease, and global climate change. Some of the characteristics of amphibians that make them useful models for studying these environmental problems are highlighted, including their trophic importance, environmental sensitivity, research tractability, and impending extinction. The article provides specific examples from the recent literature to illustrate how studies on amphibians have been instrumental in guiding scientific thought on a broad scale. Included are examples of how amphibian research has transformed scientific disciplines, generated new theories about global health, called into question widely accepted scientific paradigms, and raised awareness in the general public that our daily actions may have widespread repercussions. In addition, studies on amphibian declines have provided insight into the complexity in which multiple independent factors may interact with one another to produce catastrophic and sometimes unpredictable effects. Because of the complexity of these problems, amphibian ecologists have been among the strongest advocates for interdisciplinary research. Future studies of amphibians will be important not only for their conservation but also for the conservation of other species, critical habitats, and entire ecosystems.     

Evolutionary causes and consequences of sequential polyandry in anuran amphibians

Among anuran amphibians (frogs and toads), there are two types of polyandry: simultaneous polyandry, where sperm from multiple males compete to fertilize eggs, and sequential polyandry, where eggs from a single female are fertilized by multiple males in a series of temporally separate mating events, and sperm competition is absent. Here we review the occurrence of sequential polyandry in anuran amphibians, outline theoretical explanations for the evolution of this mating system and discuss potential evolutionary implications. Sequential polyandry has been reported in a limited number of anurans, but its widespread taxonomic and geographic distribution suggests it may be common. There have been no empirical studies that have explicitly investigated the evolutionary consequences of sequential polyandry in anurans, but species with this mating pattern share an array of behavioural, morphological and physiological characteristics, suggesting that there has been common sexual selection on their reproductive system. Sequential polyandry may have a number of adaptive benefits, including spreading the risk of brood failure in unpredictable environments, insuring against male infertility, or providing genetic benefits, either through good genes, intrinsic compatibility or genetic diversity effects. Anurans with sequential polyandry provide untapped opportunities for innovative research approaches that will contribute significantly to understanding anuran evolution and also, more broadly, to the development of sexual‐selection and life‐history theory.     

The assessment of sublethal effects of pollutants in the sea. Review of the problems.

Sublethal effects of pollution may be significant to survival of a stock of marine fish or even a species. Such effects sometimes lead to reproductive failure and have been identified so far only in freshwater systems. Atlantic salmon have disappeared from many streams in Europe and eastern North America, partly as a result of pollution in their freshwater spawning areas and in their estuarine nursing grounds. Reductions in populations of marine fishes due to pollution solely have not yet been demonstrated. However, Baltic Sea seals, where reproductive failure is apparently associated with high concentrations of DDT and polychlorinated biphenyl in the blubber, may have suffered a decline owing to the presence of these organochlorines. Sublethal effects of pollutants have been studied in the laboratory, essentially under four categories: (1) physiology (growth, swimming performance, respiration, circulation); (2) biochemistry/cell structure (blood chemistry, enzyme activity, endocrinology, histochemistry); (3) behaviour/neurophysiology; and (4) reproduction. Not all pollutants elicit meaningful responses in all categories, and a response is not always linear with pollutant concentration. For application to survival of populations the response has to be ultimately related to a healthy progression through a full life cycle, including successful reproduction. In recent time, physiological studies have moved into polluted marine environments with mobile laboratories having continuous sampling capability, to observe effects of pollutants in situ on marine organisms. The Controlled Ecosystem Pollution Experiment (Cepex) in Saanich Inlet, British Columbia, endeavours to investigate the effects of low concentrations of pollutants on marine organisms in large plastic silos having a slow replacement of water.     

水生生态环境中捕食信息素的生态学效应

捕食信息素是捕食者释放的,能够引发猎物反捕食反应的化学信号。在水生生态系统中,捕食信息素在捕食者和猎物之间信息传递及协同进化过程中发挥着重要的作用,其生态学效应在国际上受到广泛关注。捕食信息素的来源有多种形式,研究中常使用养殖过捕食者的水溶液作为捕食信息素的来源。捕食信息素的作用效果受到捕食者和猎物的种类、信息素的浓度、观察的指标等多方面因素的影响。捕食信息素可以对水生生物的行为、形态和生活史特征等方面造成影响。水生生物通过感知捕食信息素来提前预知潜在的被捕食风险,并作出适应性调整,以降低被捕食的风险。在某些情况下,捕食信息素可以与污染物产生交互作用,从而干扰污染物对水生生物的毒性。对水生环境中捕食信息素的研究现状做了综述,介绍了当前对捕食信息素来源和理化性质等本质问题的认识,总结捕食信息素对水生生物行为、形态和生活史特征的影响,以及捕食信息素对污染物毒性的干扰,并分析了这一研究领域尚存在的困难和今后的研究方向。加强对捕食信息素的研究,将为解析水生环境中捕食者和猎物的生态关系提供新依据。     

Amphibian Temperature Regulation Studies in the Field and Laboratory

Studies on thermoregulation in the laboratory and field havecome a long way from the early work done between 1940 and 1960.While some physiological studies on amphibians have progressedat the same rate as those on reptiles, field studies have beenfar behind. Laboratory studies have largely delt with thermalacclimation, evaporative water loss, and thermal and moisturegradient behavior. Field studies, following early summariesof body temperatures of field animals, have stressed behavioralthermoregulation; yet, detailed studies on behavioral thermoregulationin amphibians have been completed for only a handful of species.A few studies have placed behavioral and physiological thermoregulationinto an ecological or energetic framework; these studies arereviewed, and suggestions are made for future work.     

A review on emerging persistent organic pollutants: Current scenario in Pakistan

During the last few years, a new drift on screening of persistent organic pollutants (POPs) present in the environment of Pakistan has been observed. However, across the globe a number of reports have been devoted to the screening levels, distribution, and risk assessment and on the emission of POPs. In the case of Pakistan, the knowledge achieved and understanding of POPs contamination in the environmental compartments are still limited. Recently published literature has been a key to explore the mystery of new emerging POPs from the environment of the country. In this review, an effort was made to summarize the results of recently published reports on POPs (PCNs, PBDEs, DPs, and PCBs) from biotic and abiotic environments of Pakistan. This review also presents the available data published to date for organochlorines. The results of previously reported studies reflected that newly emerging POPs were influenced by the industrial and urban fractions and were in line with the distribution pattern of other regions of the world. These results revealed that urgent attention must be paid to these new emerging POPs, as they are reported to be present in considerable concentrations. Such detected concentrations of these banned pollutants should be checked/screened by scientific authorities to avoid adverse health risks to humans and animals.     

An examination of amphibian sensitivity to environmental contaminants: are amphibians poor canaries?

Nearly two decades ago, the global biodiversity crisis was catapulted to the front pages of newspapers with the recognition of worldwide amphibian declines. Amphibians earned their appellation, 'canaries in a coal mine', because of apparent high sensitivity to human-mediated environmental change. The most frequently cited causes for high susceptibility include permeable skin, a dual aquatic-terrestrial life cycle and a relatively rudimentary immune system. While some researchers have questioned the basis for the canary assertion, there has been no systematic evaluation of amphibian sensitivity to environmental challenges relative to other taxa. Here, we apply a database representing thousands of toxicity tests to compare the responses of amphibians relative to that of other taxonomic groups. The use of standardized methods combined with large numbers of identical challenges enables a particularly powerful test of relative effect size. Overall, we found that amphibians only exhibit moderate relative responses to water-borne toxins. Our findings imply that, as far as chemical contaminants are concerned, amphibians are not particularly sensitive and might more aptly be described as 'miners in a coal mine'. To the extent that amphibian declines have been mediated by chemical contaminants, our findings suggest that population losses and extinctions may have already occurred in a variety of taxa much more sensitive than amphibians.
Ecology Letters (2010) 13: 60–67     

Testing for Effects of UV‐B Radiation on Anti‐Predator Behavior in Amphibians: A Critique and Some Suggestions

In a recent paper, Kats et al. (2000) reported three experiments intended to test for effects of UV‐B radiation (hereafter, UV‐B) on predator‐avoidance behaviors in three amphibian species. They introduced their study in the context of concern about increasing penetration of solar UV‐B to ground level, owing to loss of stratospheric ozone, and its possible effects on amphibian populations. Kats et al. concluded from their results: `ultraviolet exposure may have important sub‐lethal effects in amphibians that could adversely effect (sic) their fitness'. As Kats et al. cited field studies that apparently demonstrated lethal effects of ambient levels of UV‐B on amphibians, their conclusion that sub‐lethal effects may occur is not obviously contentious. Furthermore, their experiments appear to be based on accepted methods. Nevertheless, I contend that the conclusions drawn by Kats et al. are not justified by the results they reported. Here, I discuss some problems with their study and suggest some possible improvements for future studies.     

The Value of Well-Designed Experiments in Studying Diseases with Special Reference to Amphibians

Relatively few studies of amphibian diseases have employed standard ecological experimental designs. We discuss what constitutes a well-designed ecological experiment and encourage their use in disease studies. We illustrate how well-designed experiments can be used to determine the effects of pathogens on amphibians and we illustrate how ancillary information, including that collected using molecular tools, can be used to enhance the value of such experiments.     

Ecology of helminth communities in tropical Australian amphibians.

Less than 50% of Australian amphibians have been recorded as hosts for helminth parasites. Despite developments in parasite community ecology in amphibians elsewhere, Australia lags behind with only two publications on this subject. Reasons advanced for this are that much of the collecting and taxonomic studies were conducted earlier this century before more recent discoveries of host genera and species as well as species complexes in the amphibian fauna. Consequently, there is a need for re-collection of hosts and parasites, and taxonomic revision of the parasites. In addition, as shown in this study, the parasite fauna in Australian amphibians is depauperate. Composition of the parasite fauna was largely dependent on the ecological associations of the host animal species. Frogs were infected with few helminth species and these occurred at low intensity, indicating, as in Europe and North America, that a depauperate fauna is also characteristic of amphibians in tropical regions.     

Global stressors and the global decline of amphibians: tipping the stress immunocompetency axis

There is a widespread consensus that the earth is experiencing a mass extinction event and at the forefront are amphibians, the most threatened of all vertebrate taxa. A recent assessment found that nearly one-third (32%, 1,856 species) of the world’s amphibian species are threatened. Amphibians have existed on the earth for over 300 million years, yet in just the last two decades there have been an alarming number of extinctions, nearly 168 species are believed to have gone extinct and at least 2,469 (43%) more have populations that are declining. Infectious diseases have been recognized as one major cause of worldwide amphibian population declines. This could be the result of the appearance of novel pathogens, or it could be that exposure to environmental stressors is increasing the susceptibility of amphibians to opportunistic pathogens. Here I review the potential effects of stressors on disease susceptibility in amphibians and relate this to disease emergence in human and other wildlife populations. I will present a series of case studies that illustrate the role of stress in disease outbreaks that have resulted in amphibian declines. First, I will examine how elevated sea-surface temperatures in the tropical Pacific since the mid-1970s have affected climate over much of the world and could be setting the stage for pathogen-mediated amphibian declines in many regions. Finally, I will discuss how the apparently rapid increase in the prevalence of amphibian limb deformities is linked to the synergistic effects of trematode infection and exposure to chemical contaminants.     

Batrachology of Japan and adjacent regions - a systematic review

As is the case with most other organisms, studies of amphibians started with systematics as the basis for many other biological fields, but systematics itself has endured a slower progress than other fields of biology have enjoyed for a long time. Nevertheless, philosophical and methodological 'revolutions' in the past 30 years, combined with more recent progress in biochemical and molecular techniques, have been forcing current systematics into a great change. Stejneger's (1907) monumental book, the first thorough review of amphibians from Japan and adjacent regions, was published nearly a century ago, and since then, the increase in number of areas that have been closely surveyed, together with the application of new theories and methods, have greatly enlarged our knowledge of the amphibian fauna in this region. For example, extensive experiments by the Hiroshima School using artificial hybridization techniques greatly contributed to our understandings of relationships among some groups of amphibians that had been taxonomically confusing. In addition to this, new species discovered by more detailed field surveys in a wider area and cryptic species revealed by analyses of isozymes and acoustic properties have made for a great increment in the number of taxa in this region. Further, analyses of DNA sequences now enable us to infer the affinities of taxa with few phylogenetically informative phenotypic properties, and the resultant phylogenetic hypotheses further contribute to our understandings of biogeography and many other issues on amphibian evolution. From these advances, it has been clarified that the amphibian species diversity is far greater than was previously expected, which in turn means a high endemism within particular regions. It is expected that future studies will also reveal exact systematic relationships of wide-ranging species, both those occurring within this region and those extending outside of it. For that purpose, international and interdisciplinary studies are indispensable. But what is also necessary is to conserve amphibian populations so that they will not disappear before their true relationships are realized. Knowledge of systematics among young amphibian biologists is also a matter of high priority.     

Chemosignals,hormones, and amphibian reproduction

This article is part of a Special Issue “Chemosignals and Reproduction”.Amphibians are often thought of as relatively simple animals especially when compared to mammals. Yet the chemosignaling systems used by amphibians are varied and complex. Amphibian chemosignals are particularly important in reproduction, in both aquatic and terrestrial environments. Chemosignaling is most evident in salamanders and newts, but increasing evidence indicates that chemical communication facilitates reproduction in frogs and toads as well. Reproductive hormones shape the production, dissemination, detection, and responsiveness to chemosignals. A large variety of chemosignals have been identified, ranging from simple, invariant chemosignals to complex, variable blends of chemosignals. Although some chemosignals elicit straightforward responses, others have relatively subtle effects. Review of amphibian chemosignaling reveals a number of issues to be resolved, including: 1) the significance of the complex, individually variable blends of courtship chemosignals found in some salamanders, 2) the behavioral and/or physiological functions of chemosignals found in anuran “breeding glands”, 3) the ligands for amphibian V2Rs, especially V2Rs expressed in the main olfactory epithelium, and 4) the mechanism whereby transdermal delivery of chemosignals influences behavior. To date, only a handful of the more than 7000 species of amphibians has been examined. Further study of amphibians should provide additional insight to the role of chemosignals in reproduction.     

Global tests of regional effect on species richness of vascular plants and terrestrial vertebrates

Determining the effects of regional and contemporary factors on large-scale patterns in species richness has been a fundamental question in modern ecology and biogeography. However, few studies have examined effects of historical and regional factors on species richness at the global scale, and conclusions are often inconsistent or controversial. Here, I use a comprehensive dataset to examine regional effects on species richness of vascular plants and four taxa of terrestrial vertebrates (mammals, birds, reptiles, and amphibians) in the same set of sample units (ecoregions) in seven biogeographic realms across the globe. The same spatial scale and the same set of environmental variables, which are thought to influence large-scale patterns in species richness of vascular plants and terrestrial vertebrates, are used for all the five taxa. Species richness of each taxon is compared across biogeographic realms. Regional effect on species richness has been found for all the five taxa. Of the 90 realm-pair comparisons for the five taxa between observed richness of a region and the richness of the region predicted by the richness–environment relationship derived from the data of another region, 74 (82.2%) showed significant differences between observed and predicted species richness, indicating that a species richness–environment relationship developed for one region cannot accurately predict species richness in other regions of similar environments.     

Grazing refuges, external avoidance of herbivory and plant diversity

Avoidance and tolerance are the two means by which plants cope with herbivores. Avoidances internal to the plant, such as morphology, chemical repellants, thorns, etc., have received considerable attention in the plant‐herbivore literature, but relatively little consideration has been given to avoidances external to the plant. We develop a conceptual framework of external plant avoidances of herbivory based on foraging selection impedances (associational avoidances), behavioral impedances (indirect avoidances), and physical impedances (refuges) organized along axes of efficiency, degree of protection, and necessity of tolerance characteristics. Associational avoidances are uncommon for terrestrial mammalian herbivores compared to plant‐insect or marine situations. Indirect avoidances mediated through herbivore territoriality, predator avoidance, and other behaviors independent of foraging decisions are probably common in nature, but few have been formally documented. Biotic and geologic refuges providing a physical impedance are the only avoidances shown to have implications for plant biodiversity. This is particularly true for geologic refuges, where there is not a tradeoff between competition and the refuge effect. Small geologic refuges (rock outcrops, cliffs, etc.) are more likely to also positively or negatively alter associated plant microenvironments than large geologic refuges (mesas, islands, etc.). In a survey, 86% of small refuge studies reported positive effects on plant diversity compared to 50% for large refuges. Geologic refuges in more productive environments were more important in protecting diversity than refuges in less productive, semiarid environments, and the effects of protection were greater in communities with short compared to long evolutionary histories of grazing. Other characteristics of refuges such as extent across the landscape and the manner they alter or ameliorate the environment, as well as characteristics of the herbivore such as small or large, generalist or specialist may also determine the effectiveness of refuges, but there are too few studies to assess these factors.