Widespread occurrence of ranavirus in pond-breeding amphibian populations

Ranaviruses are an emerging threat for many amphibian populations, yet their distribution in amphibian communities and the association of infection with possible stressors and species is not fully understood due to historically sparse surveillance. Agricultural practices that reduce the water quality of amphibian breeding habitats (e.g., cattle access to wetlands) and environmental stressors (e.g., lower temperatures) may contribute to ranavirus emergence. We tested larval amphibians for ranavirus infection across four seasons in farm ponds (n?=?40) located in Tennessee, USA. Cattle at various densities were allowed access to half of the sampled ponds. Ranavirus infections were detected in nine species and in 33 of the sampled ponds (83%), illustrating widespread occurrence of the pathogen. Species within the family Ranidae were the most frequently infected. In 13 of the ponds containing infected individuals, prevalence exceeded 40% during at least one season. Infections were detected in multiple seasons in 20 of the sampled ponds containing infections, suggesting that ranaviruses are relatively persistent in these systems. Cattle had negative effects on water quality (turbidity and ammonia) and there was a positive association between cattle abundance and ranavirus prevalence in the summer. Counter to previous field studies in North America, we found a significant positive association between water temperature and ranavirus prevalence in the fall sampling events. Despite these findings, the influences of cattle and temperature on ranavirus prevalence were not consistent across seasons. As such, the mechanisms driving high ranavirus prevalence across the landscape and over time remain unclear. Given the widespread occurrence of ranaviruses in wild amphibians, we encourage the implementation of surveillance programs to help identify potential drivers of emergence. Sites with high ranavirus prevalence should be monitored annually for outbreaks, and the long-term effects on population size determined.     

Spatial distribution of colonizing Listronotus maculicollis populations: implications for targeted management and host preference

We characterized the spatio‐temporal distribution of emerging overwintered adult Listronotus maculicollis populations colonizing golf course fairways with Spatial Analyses by Distance IndicEs (SADIE) with the goal to better target management tactics and to test assumptions for weevil preference for annual bluegrass (Poa annua). Adults randomly colonized and moved throughout fairways. However, cumulative captures were significantly aggregated along fairway edges closest to overwintering sites demonstrating progressive movement through the edges. Spatial association analyses suggest that the spatial patterns of cumulative captures of adults rather than weekly patterns were strongly associated with larvae, indicating that the adults enter fairways from the edges and deposit eggs over the course of several weeks. We did not observe an effect of host species on the distribution of either adult or larval L. maculicollis which leads us to question traditional assumptions of host preference for short mown P. annua. Instead, we propose that the aggregated distribution of larvae is generated by a low encounter rate of short mown hosts, rather than preference for species or cultivar. This study indicates that caution need to be applied when using preference‐performance criteria in host preference studies. Future behavioural studies need to address the contributions of encounter rate and host species on L. maculicollis host selection and oviposition.     

Network-scale effects of invasive species on spatially-structured amphibian populations

Understanding the factors affecting the dynamics of spatially-structured populations (SSP) is a central topic of conservation and landscape ecology. Invasive alien species are increasingly important drivers of the dynamics of native species. However, the impacts of invasives are often assessed at the patch scale, while their effects on SSP dynamics are rarely considered. We used long-term abundance data to test whether the impact of invasive crayfish on subpopulations can also affect the whole SSP dynamics, through their influence on source populations. From 2010 to 2018, we surveyed a network of 58 ponds and recorded the abundance of Italian agile frog clutches, the occurrence of an invasive crayfish, and environmental features. Using Bayesian hierarchical models, we assessed relationhips between frog abundance in ponds and a) environmental features; b) connectivity within the SSP; c) occurrence of invasive species at both the patch- and the SSP-levels. If spatial relationships between ponds were overlooked, we did not detect effects of crayfish presence on frog abundance or trends. When we jointly considered habitat, subpopulation and SSP features, processes acting at all these levels affected frog abundance. At the subpopulation scale, frog abundance in a year was related to habitat features, but was unrelated to crayfish occurrence at that site during the previous year. However, when we considered the SSP level, we found a strong negative relationship between frog abundance in a given site and crayfish frequency in surrounding wetlands during the previous year. Hence, SSP-level analyses can identify effects that would remain unnoticed when focussing on single patches. Invasive species can affect population dynamics even in not invaded patches, through the degradation of subpopulation networks. Patch-scale assessments of the impact of invasive species can thus be insufficient: predicting the long-term interplay between invasive and native populations requires landscape-level approaches accounting for the complexity of spatial interactions.     

Dispersal and the metapopulation paradigm in amphibian ecology and conservation: are all amphibian populations metapopulations?

Amphibians are frequently characterized as having limited dispersal abilities, strong site fidelity and spatially disjunct breeding habitat. As such, pond‐breeding species are often alleged to form metapopulations. Amphibian species worldwide appear to be suffering population level declines caused, at least in part, by the degradation and fragmentation of habitat and the intervening areas between habitat patches. If the simplification of amphibians occupying metapopulations is accurate, then a regionally based conservation strategy, informed by metapopulation theory, is a powerful tool to estimate the isolation and extinction risk of ponds or populations. However, to date no attempt to assess the class‐wide generalization of amphibian populations as metapopulations has been made. We reviewed the literature on amphibians as metapopulations (53 journal articles or theses) and amphibian dispersal (166 journal articles or theses for 53 anuran species and 37 salamander species) to evaluate whether the conditions for metapopulation structure had been tested, whether pond isolation was based only on the assumption of limited dispersal, and whether amphibian dispersal was uniformly limited. We found that in the majority of cases (74%) the assumptions of the metapopulation paradigm were not tested. Breeding patch isolation via limited dispersal and/or strong site fidelity was the most frequently implicated or tested metapopulation condition, however we found strong evidence that amphibian dispersal is not as uniformly limited as is often thought. The frequency distribution of maximum movements for anurans and salamanders was well described by an inverse power law. This relationship predicts that distances beneath 11–13 and 8–9 km, respectively, are in a range that they may receive one emigrating individual. Populations isolated by distances approaching this range are perhaps more likely to exhibit metapopulation structure than less isolated populations. Those studies that covered larger areas also tended to report longer maximum movement distances – a pattern with implications for the design of mark‐recapture studies. Caution should be exercised in the application of the metapopulation approach to amphibian population conservation. Some amphibian populations are structured as metapopulations – but not all.     

Complexity in conservation: lessons from the global decline of amphibian populations

As part of an overall "biodiversity crisis" many amphibian populations are in decline throughout the world. Numerous causes have been invoked to explain these declines. These include habitat destruction, climate change, increasing levels of ultraviolet radiation, environmental contamination, disease, and the introduction of non-native species. In this paper, we argue that amphibian population declines are caused by different abiotic and biotic factors acting together in a context-dependent fashion. Moreover, different species and different populations of the same species may react in different ways to the same environmental insult. Thus, the causes of amphibian population declines will vary spatially and temporally. Although some generalizations (e.g. those concerning environmental stress and disease outbreaks) can be made about amphibian population declines, we suggest that these generalizations take into account the context-dependent dynamics of ecological systems.     

Analysis of three amphibian populations with quarter-century long time-series.

Amphibians are in decline in many parts of the world. Long tme-series of amphibian populations are necessary to distinguish declines from the often strong fluctuations observed in natural populations. Time-series may also help to understand the causes of these declines. We analysed 23-28-year long time-series of the frog Rana temporaria. Only one of the three studied populations showed a negative trend which was probably caused by the introduction of fish. Two populations appeared to be density regulated. Rainfall had no obvious effect on the population fluctuations. Whereas long-term studies of amphibian populations are valuable to document population declines, most are too short to reveal those factors that govern population dynamics or cause amphibian populations to decline.     

Non‐native salmonids affect amphibian occupancy at multiple spatial scales

Aim The introduction of non‐native species into aquatic environments has been linked with local extinctions and altered distributions of native species. We investigated the effect of non‐native salmonids on the occupancy of two native amphibians, the long‐toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), across three spatial scales: water bodies, small catchments and large catchments. Location Mountain lakes at ≥ 1500 m elevation were surveyed across the northern Rocky Mountains, USA. Methods We surveyed 2267 water bodies for amphibian occupancy (based on evidence of reproduction) and fish presence between 1986 and 2002 and modelled the probability of amphibian occupancy at each spatial scale in relation to habitat availability and quality and fish presence. Results After accounting for habitat features, we estimated that A. macrodactylum was 2.3 times more likely to breed in fishless water bodies than in water bodies with fish. Ambystoma macrodactylum also was more likely to occupy small catchments where none of the water bodies contained fish than in catchments where at least one water body contained fish. However, the probability of salamander occupancy in small catchments was also influenced by habitat availability (i.e. the number of water bodies within a catchment) and suitability of remaining fishless water bodies. We found no relationship between fish presence and salamander occupancy at the large‐catchment scale, probably because of increased habitat availability. In contrast to A. macrodactylum, we found no relationship between fish presence and R. luteiventris occupancy at any scale. Main conclusions Our results suggest that the negative effects of non‐native salmonids can extend beyond the boundaries of individual water bodies and increase A. macrodactylum extinction risk at landscape scales. We suspect that niche overlap between non‐native fish and A. macrodactylum at higher elevations in the northern Rocky Mountains may lead to extinction in catchments with limited suitable habitat.     

Neotropical tadpoles influence stream benthos: evidence for the ecological consequences of decline in amphibian populations

1. Few studies have assessed the role of tadpoles in tropical streams, although they are often abundant and conspicuous components of these systems. Moreover, amphibian populations are declining around the globe, particularly stream‐dwelling species in tropical uplands, and the ecological consequences of these losses are not understood. 2. We chose a stream in the central Panamanian highlands, which has an intact fauna of stream‐breeding anurans, to examine the ecological consequences of amphibian losses. This site differs dramatically from sites in nearby western Panama and Costa Rica where anuran diversity and abundance have declined greatly in the last two decades. 3. We used an underwater electric field to create tadpole exclosures in runs, so that we could evaluate their influence on sediment dynamics and the abundance and community structure of algae and aquatic insects. Tadpoles reduced total sediments and both organic and inorganic fractions on substrata. Tadpoles also reduced algal abundance on substrata by approximately 50% and decreased algal biovolume. Gut content analyses showed that tadpoles consumed algae and sediments and we could see that algae and sediments were also displaced through bioturbation. 4. Atelopus zeteki, Rana warszewitschii, and Hyla spp. were the dominant larval anurans responsible for the effects observed. Visual surveys indicated that the densities of these taxa ranged from 23 (R. warszewitschii and Hyla spp. combined) to 43 m^?2 (A. zeteki) in runs. 5. The abundance of baetid mayflies was lower in tadpole exclosures compared with controls, and this was attributed to tadpoles facilitating mayfly feeding by removing sediments and exposing underlying periphyton. 6. Tadpoles affect the abundance and diversity of basal resources and other primary consumers, and thus influence food web dynamics and energy flow in these tropical streams. Catastrophic decline in stream‐breeding anuran populations will influence the structure and function of neotropical stream ecosystems.     

Spatial distribution of competing populations

We consider spatial distributions of two competing and diffusing populations whose habitats are partly overlapping. As a model, certain reaction—diffusion equations are used in the finite and in the infinite regions with Dirichlet boundary conditions. On the assumption of extremely different diffusive rates of the two species, it is verified, by the use of singular perturbation techniques, that the slowly diffusing species can survive in some subregions, although the species with the greater diffusive rate rapidly occupies the region at the initial stage, and that coexistence of two populations is realized, reducing the effect of interspecific competition by spatial segregation. It will be also shown that the size of the region where the slowly moving population can survive exhibits a markedly qualitative change, depending on the values of some parameters, and that the population can extends the distribution infinitely, when the parameters satisfy a certain condition.     

Spatial network structure and amphibian persistence in stochastic environments

In the past few years, the framework of complex networks has provided new insight into the organization and function of biological systems. However, in spite of its potential, spatial ecology has not yet fully incorporated tools and concepts from network theory. In the present study, we identify a large spatial network of temporary ponds, which are used as breeding sites by several amphibian species. We investigate how the structural properties of the spatial network change as a function of the amphibian dispersal distance and the hydric conditions. Our measures of network topology suggest that the observed spatial structure of ponds is robust to drought (compared with similar random structures), allowing the movement of amphibians to and between flooded ponds, and hence, increasing the probability of reproduction even in dry seasons.     

Current threats faced by amphibian populations in the southern cone of South America

In this work, we update and increase knowledge on the severity and extent of threats affecting 57 populations of 46 amphibian species from Chile and Argentina in southern South America. We analyzed the intrinsic conservation problems that directly impact these populations. We shared a questionnaire among specialists on threats affecting target amphibian populations with information on i) range, ii) historical occurrence and abundance, iii) population trends, iv) local extinctions, v) threats, and vi) ongoing and necessary conservation/research. We assessed association patterns between reported threats and population trends using multiple correspondence analysis. Since 2010, 25 of 57 populations have declined, while 16 experienced local extinctions. These populations were affected by 81% of the threat categories analyzed, with those related to agricultural activities and/or habitat modifications being the most frequently reported. Invasive species, emerging diseases, and activities related to grazing, ranching, or farming were the threats most associated with population declines. Low connectivity was the most frequent intrinsic conservation problem affecting 68% of the target populations, followed by low population numbers, affecting 60%. Ongoing monitoring activity was conducted in 32 (56%) populations and was the most frequent research activity. Threat mitigation was reported in 27 (47%) populations and was the most frequent ongoing management activity. We found that habitat management is ongoing in 5 (9%) populations. At least 44% of the amphibian populations surveyed in Chile and Argentina are declining. More information related to the effect of management actions to restore habitats, recover populations, and eliminate threats such as invasive species is urgently needed to reverse the conservation crisis facing amphibians in this Neotropical region.     

Multiple zoosporic parasites pose a significant threat to amphibian populations

There is substantial evidence for the dominant role of Batrachochytrium dendrobatidis in amphibian population dynamics. However, a wide range of other pathogens could also be important in precipitating amphibian population declines, particularly in the face of climate change or other stressors. Here we discuss some examples of zoosporic parasites in the Chytridiomycota, Mesomycetozoa, Perkinsozoa and Oomycota, all of which infect amphibians in freshwater habitats. The pathosystem model provides an excellent basis for understanding host–parasite interactions. Chemotactic zoopores and several families of proteases facilitate infection. Introduction of non-native host may accelerate the dispersal of these parasites. Unlike B. dendrobatidis some of the other zoosporic parasites grow well at or slightly above 25 °C, and their growth rates are likely to increase with global warming. The interactions of parasites with each other and the combined effect of simultaneous infection with multiple species in amphibian populations remain to be carefully studied.     

Observing micro-evolutionary processes of viral populations at multiple scales

Advances in sequencing technology coupled with new integrative approaches to data analysis provide a potentially transformative opportunity to use pathogen genome data to advance our understanding of transmission. However, to maximize the insights such genetic data can provide, we need to understand more about how the microevolution of pathogens is observed at different scales of biological organization. Here, we examine the evolutionary processes in foot-and-mouth disease virus observed at different scales, ranging from the tissue, animal, herd and region. At each scale, we observe analogous processes of population expansion, mutation and selection resulting in the accumulation of mutations over increasing time scales. While the current data are limited, rates of nucleotide substitution appear to be faster over individual-to-individual transmission events compared with those observed at a within-individual scale suggesting that viral population bottlenecks between individuals facilitate the fixation of polymorphisms. Longer-term rates of nucleotide substitution were found to be equivalent in individual-to-individual transmission compared with herd-to-herd transmission indicating that viral diversification at the herd level is not retained at a regional scale.     

An insight into the skin glands,dermal scales and secretions of the caecilian amphibian Ichthyophis beddomei

The caecilian amphibians are richly endowed with cutaneous glands, which produce secretory materials that facilitate survival in the hostile subterranean environment. Although India has a fairly abundant distribution of caecilians, there are only very few studies on their skin and secretion. In this background, the skin of Ichthyophis beddomei from the Western Ghats of Kerala, India, was subjected to light and electron microscopic analyses. There are two types of dermal glands, mucous and granular. The mucous gland has a lumen, which is packed with a mucous. The mucous-producing cells are located around the lumen. In the granular gland, a lumen is absent; the bloated secretory cells, filling the gland, are densely packed with granules of different sizes which are elegantly revealed in TEM. There is a lining of myo-epithelial cells in the peripheral regions of the glands. Small flat disk-like dermal scales, dense with squamulae, are embedded in pockets in the dermis, distributed among the cutaneous glands. 1–4 scales of various sizes are present in each scale pocket. Scanning electron microscopic observation of the skin surface revealed numerous glandular openings. The skin gland secretions, exuded through the pores, contain fatty acids, alcohols, steroid, hydrocarbons, terpene, aldehyde and a few unknown compounds.     

Identifying monitoring gaps for amphibian populations in a North American biodiversity hotspot,the southeastern USA

I review the primary literature to ascertain the status of amphibian monitoring efforts in the southeastern USA, a “hotspot” for biodiversity in North America. This effort revealed taxonomic, geographic and ecological disparities in studies of amphibian populations in this region. Of the species of anurans and caudates known to occur in the Southeast, 73.8 and 33.3 %, respectively, have been monitored continuously for at least 4 years. Anurans are generally shorter-lived than are caudates and, thus, have been studied for the equivalent of at least one population turnover more than have caudates. The percentage of species (of those occurring in a given state) monitored continuously for at least 4 years was lowest for Alabama and Mississippi and highest for Florida for both taxa. The vast majority of studies (69.6 %) were conducted on species that inhabit natural freshwater wetlands, in contrast to other aquatic and terrestrial habitats. Species considered threatened by the International Union for Conservation of Nature comprised only 7.7 % of 65 species that have been studied consistently. The majority of comparative studies of contemporary versus historical occurrences were potentially biased by the use of “presence-only” historical data and resurveys of short duration. Other issues, such as inadequate temporal and spatial scale and neglect of different sources of error, were common. Awareness of these data gaps and sampling and statistical issues may help facilitate informed decisions in setting future monitoring priorities, particularly with respect to species, habitats and locations that have been largely overlooked in past and ongoing studies.