Chytrid fungus infection in alpine tree frogs is associated with individual heterozygosity and population isolation but not population-genetic diversity

Chytridiomycosis, a disease caused by the emerging fungus Batrachochytrium dendrobatidis (Bd), has been implicated in the decline of over 500 amphibian species. Population declines could have important genetic consequences, including reduced genetic diversity. We contrasted genetic diversity among both long-Bd-exposed and unexposed populations of the south-east Australian alpine tree frog (Litoria verreauxii alpina) across its range. At the population level, we found no significant differences in genetic diversity between Bd-exposed and unexposed populations. Encouragingly, even Bd-infected remnant populations that are now highly isolated maintain genetic diversity comparable to populations in which Bd is absent. Spatial genetic structure among populations followed an isolation-by-distance pattern, suggesting restricted movement among remnant populations. At the individual level, greater heterozygosity was associated with reduced probability of infection. Loss of genetic diversity in remnant populations that survived chytridiomycosis epidemics does not appear to be a threat to L. v. alpina. We suggest several factors underpinning maintenance of genetic diversity: (1) remnant populations have remained large enough to avoid losses of genetic diversity; (2) many individuals in the population are able to breed once before succumbing to disease; and (3) juveniles in the terrestrial environment have low exposure to Bd, providing an annual ‘reservoir’ of genetic diversity. The association between individual heterozygosity and infection status suggests that, while other work has shown all breeding adults are typically killed by Bd, males with greater heterozygosity may survive longer and obtain fitness benefits through extended breeding opportunities. Our results highlight the critical role of life history in mitigating the impacts of Bd infection for some amphibian species, but we infer that increased isolation as a result of disease-induced population extirpations will enhance population differentiation and thus biogeographic structure

Comparative use of active searches and artificial refuges to detect amphibians in terrestrial environments

Artificial refuges (cover boards) are commonly used to survey and monitor herpetofauna in many parts of the world. Despite the extensive use of artificial refuges in mesic environments, their effectiveness for detecting amphibians in temperate zones has rarely been examined. We compared amphibian detection probabilities between two survey methods; active searches of natural habitat and artificial refuges of three different types (corrugated steel, roofing tiles and timber railway sleepers). Our study area included five bioregions encompassing a 1,180 km latitudinal gradient across a modified, temperate eucalypt woodland vegetation community in south-eastern Australia. We deployed 14,778 artificial refuges in terrestrial environments, within patches of remnant vegetation, and collected presence and abundance data on herpetofauna between 1999 and 2017. We used Bayesian logistic regression to identify the most effective survey method for detecting frog species across all bioregions. We modelled frog detections by fitting survey method, time since refuge deployment, and rainfall prior to each survey. We detected 3970 individuals from 18 frog species. Overall, we found active searches and timber substrates most effective for detecting a broad range of species, although detection rates were driven by the numerically abundant spotted marsh frog Limnodynastes tasmaniensis. Timber refuges were effective for detecting several burrowing species, whereas active searches were effective at detecting habitat generalists. Quadratic effects of rainfall prior to survey as opposed to linear effects of time since artificial refuge placement was important in explaining frog detection rates in some bioregions. Active searches, timber railway sleepers and sheets of corrugated steel provide complimentary survey methods for detecting amphibians, although detection rates are influenced by rainfall patterns. Artificial refuges provide a time-effective and standardised method for studying amphibians in their non-breeding terrestrial environment and should be incorporated into future surveys and biodiversity monitoring programs.The work was funded by the Australian Govern-ment's Caring for Our Country Initiative, the National Environmental Science Program's Threatened Species Recovery Hub and the Australian Research Council, and is supported by the Murray Local Land Services, Riverina Local land Services, Central Tablelands Local land Services, North East Catchment Management Authority, Goulburn Broken Catchment Management Authority, NSW Environment Trust, Ian Potter Foundation and the Vincent Fairfax Family Foundation

Tests of predictions associated with temporal changes in Australian bird populations

Global biodiversity loss is the cumulative result of local species declines. To combat biodiversity loss, detailed information on the temporal trends of at-risk species at local scales is needed. Here we report the results of a 13-year study of temporal change in bird occupancy in one of the most heavily modified biomes worldwide; the temperate woodlands of south-eastern Australia. We sought to determine if temporal changes in bird species were different between three broad native vegetation types (old-growth woodland, regrowth woodland and restoration plantings) and between species traits (body size, migratory status, rarity, woodland dependency, or diet). We found evidence of decline for over a quarter of all bird species for which we had sufficient data for detailed analysis (30 out of 108 species). In contrast, only 14 species increased significantly. Temporal change of birds was linked to life-history attributes, with patterns often being habitat-dependent. Nectarivores and large-bodied birds declined across all vegetation types, whereas small-bodied species increased, particularly in restoration plantings. Contrasting with patterns documented elsewhere, resident but not migratory species declined, with this trend strongest in restoration plantings. Finally, our analyses showed that, as a group, common birds tended to decline whereas rare birds tended to increase, with effects for both most pronounced in restoration plantings. Our results highlight the benefit of targeted restoration planting for some species, but also demonstrate that many common species that have long-persisted in human-dominated landscapes are experiencing severe declines. © 2018 Elsevier Ltd. **Please note that there are multiple authors for this article therefore only the name of the first 5 including Federation University Australia affiliate “Philip Barton” is provided in this record*

Amphibians in agricultural landscapes: the habitat value of crop areas, linear plantings and remnant woodland patches

Mitigating the negative impacts of agriculture on amphibians requires knowledge of how different land uses affect species distribution and community composition. In the case of frogs, there is currently insufficient information on their use of terrestrial habitats in cropping landscapes to inform conservation planning. We examined how four different farmland types (linear plantings, cereal crops, grazing paddocks and woody mulch) and crop harvesting influenced amphibian abundance, richness, body condition and movement. We found the abundance of frogs was significantly higher in linear plantings compared to grazing paddocks and adjacent patches of remnant woodland vegetation. However, species richness and abundance of three individual species did not vary significantly between farmland types. For the most common frog Uperoleia laevigata, body condition was higher at the edges of the woody debris treatment (coupled with higher abundance) and lower in farmland with debris and linear plantings. The body condition of Limnodynastes tasmaniensis and L. interioris was not influenced by farmland type. Frog abundance and condition was largely unaffected by crop harvesting. However, frogs were less common after harvesting at the edges of farmland and within remnant patches. Movement patterns did not suggest mass movement out of crops after harvest, where almost half of all individuals recaptured remained within the farmland. These results suggest that some generalist frog species may have an affinity for habitats within agricultural paddocks, particularly when key habitat features like plantings are present. However, we found overall frog richness was low and did not differ between remnant patches, edges and farmland which may be an indication of habitat degradation within terrestrial habitats across the landscape. Although protection of remnant native vegetation is important, conservation strategies for the protection of amphibians will be ineffective if they do not consider the variety of land uses and the relationships of different species and their microhabitats within and outside of patches.Funding was provided by the Australian Government Research Training Program, Central Tablelands Local Land Services, NSW Environmental Trust and the Lake Cowal Foundation

Spatiotemporal effects of logging and fire on tall, wet temperate eucalypt forest birds

Forests globally are subject to disturbances such as logging and fire that create complex temporal variation in spatial patterns of forest cover and stand age. However, investigations that quantify temporal changes in biodiversity in response to multiple forms of disturbance in space and time are relatively uncommon. Over a 10-yr period, we investigated the response of bird species to spatiotemporal changes in forest cover associated with logging and wildfire in the mountain ash (Eucalyptus regnans) forests of southeastern Australia. Specifically, we examined how bird occurrence changed with shifts in the proportion of area burned or logged in a 4.5 km radius surrounding our 88 long-term field survey sites, each measuring 1 ha in size. Overall species richness was greatest in older forest patches, but declined as the amount of fire around each site increased. At the individual species level, 31 of the 37 bird species we modeled exhibited a negative response to the amount of fire in the surrounding landscape, while one species responded positively to fire. Only nine species exhibited signs of recovery in the 6 yr of surveys following the fire. Five species were more likely to be detected as the proportion of logged forest surrounding a site increased, suggesting a possible "concentration effect" with displaced birds moving into unlogged areas following harvesting of adjacent areas. We also identified relationships between the coefficients of life history attributes and spatiotemporal changes in forest cover and stand age. Large-bodied birds and migratory species were associated with landscapes subject to large amounts of fire in 2009. There were associations between old growth stands and small-bodied bird species and species that were not insectivores. Our study shows that birds in mountain ash forests are strongly associated with old growth stands and exhibit complex, time-dependent, and species-specific responses to landscape disturbance. Despite logging and fire both being high-severity perturbations, no bird species exhibited similar responses to fire and logging in the landscape surrounding our sites. Thus, species responses to one kind of landscape-scale disturbance are not readily predictable based on an understanding of the responses to another kind of (albeit superficially similar) disturbance.Threatened Species Recovery Hub of the National Environmental Science Program, Parks Victoria, and Victorian Government Department of Environment, Land, Water and Plannin

Beyond the pond: Terrestrial habitat use by frogs in a changing climate

Amphibians are proportionately over-represented in the current wave of global biodiversity loss. Disease and habitat loss are implicated in many amphibian species declines, but amphibians are also predicted to be sensitive to changes in climate, particularly changes in temperature and loss of moisture. These changes could severely impact frog use of terrestrial habitats. We used data from a long-term (2007–18) landscape-scale experiment in south-eastern Australia to test the effects of terrestrial habitat characteristics and restoration treatments on frog species. We found declines in species richness and in the occurrences of two locally-common species (Limnodynastes tasmaniensis and Uperoleia laevigata). These declines were associated with high maximum temperatures, low minimum temperatures and low rainfall. Coarse woody debris addition was associated with higher species richness and L. tasmaniensis occurrence, but this effect was not greater in times of reduced rainfall and high maximum temperatures, implying a weak ‘refugia’ effect. Frogs were positively associated with wetter sites, and this association increased with higher maximum temperatures. Our findings add to a growing body of evidence that show that prolonged periods of drought pose a key threat to frog populations and that short periods of relief from drought conditions are insufficient to allow recovery of terrestrial frog populations over the long term. Restoration efforts could include the provision of coarse woody debris and should also ensure that good quality aquatic habitat, such as drought-resistant ponds and dams, are available throughout hot and dry times as a supplement to ephemeral aquatic habitat.Funding and in-kind logistic support for this project was provided by the ACT Government as part of an Australian Research Council Linkage Grant (LP0561817; LP110100126, LP140100209). ADM was supported by an Australian Research Council Future Fellowship for part of the study (FT100100358). MY was funded by the ANU Strategic Research Fund. MJE was partly funded by the Japan Society for the Promotion of Science (JSPS Postdoctoral Research Fellowship P19084)

Reservoir-host amplification of disease impact in an endangered amphibian

Emerging wildlife pathogens are an increasing threat to biodiversity. One of the most serious wildlife diseases is chytridiomycosis, caused by the fungal pathogen, Batrachochytrium dendrobatidis (Bd), which has been documented in over 500 amphibian species. Amphibians vary greatly in their susceptibility to Bd; some species tolerate infection, whereas others experience rapid mortality. Reservoir hosts-species that carry infection while maintaining high abundance but are rarely killed by disease-can increase extinction risk in highly susceptible, sympatric species. However, whether reservoir hosts amplify Bd in declining amphibian species has not been examined. We investigated the role of reservoir hosts in the decline of the threatened northern corroboree frog (Pseudophryne pengilleyi) in an amphibian community in southeastern Australia. In the laboratory, we characterized the response of a potential reservoir host, the (nondeclining) common eastern froglet (Crinia signifera), to Bd infection. In the field, we conducted frog abundance surveys and Bd sampling for both P. pengilleyi and C. signifera. We built multinomial logistic regression models to test whether Crinia signifera and environmental factors were associated with P. pengilleyi decline. C. signifera was a reservoir host for Bd. In the laboratory, many individuals maintained intense infections (>1000 zoospore equivalents) over 12 weeks without mortality, and 79% of individuals sampled in the wild also carried infections. The presence of C. signifera at a site was strongly associated with increased Bd prevalence in sympatric P. pengilleyi. Consistent with disease amplification by a reservoir host, P. pengilleyi declined at sites with high C. signifera abundance. Our results suggest that when reservoir hosts are present, population declines of susceptible species may continue long after the initial emergence of Bd, highlighting an urgent need to assess extinction risk in remnant populations of other declined amphibian species.Funding was provided by the Australian Academy of Science and a Taronga Zoo Field Conservation Grant. B. Scheele was supported by Australian Research Council grant LP110200240 during the revisions of this manuscript

High adult mortality in disease-challenged frog populations increases vulnerability to drought

Pathogen emergence can drive major changes in host population demography, with implications for population dynamics and sensitivity to environmental fluctuations. The amphibian disease chytridiomycosis, caused by infection with the fungal pathogen Batrachochytrium dendrobatidis (Bd), is implicated in the severe decline of over 200 amphibian species. In species that have declined but not become extinct, Bd persists and can cause substantial ongoing mortality. High rates of mortality associated with Bd may drive major changes in host demography, but this process is poorly understood. Here, we compared population age structure of Bd-infected populations, Bd-free populations, and museum specimens collected prior to Bd emergence for the endangered Australian frog, Litoria verreauxii alpina (alpine tree frog). We then used population simulations to investigate how pathogen-associated demographic shifts affect the ability of populations to persist in stochastic environments. We found that Bd-infected populations have a severely truncated age structure associated with very high rates of annual adult mortality. Near-complete annual adult turnover in Bd-infected populations means that individuals breed once, compared with Bd-free populations where adults may breed across multiple years. Our simulations showed that truncated age structure erodes the capacity of populations to withstand periodic recruitment failure; a common challenge for species reproducing in uncertain environments. We document previously undescribed demographic shifts associated with a globally emerging pathogen and demonstrate how these shifts alter host ecology. Truncation of age structure associated with Bd effectively reduces host niche width, and can help explain the contraction of L. v. alpina to perennial waterbodies where the risk of drought-induced recruitment failure is low. Reduced capacity to tolerate other sources of mortality may explain variation in decline severity among other chytridiomycosis-challenged species and highlights the potential to mitigate disease impacts through minimising other sources of mortality

Refracting custom in Western Sahara's quest for statehood

This article argues that distinctions made by local actors between different legal and normative orders within a broad field of custom should receive greater analytical attention. Local distinctions within custom have sometimes been overlooked in scholarly emphasis on other distinctions, such as between custom and state law, or between custom and religious law. The significance of local distinctions within custom comes to the fore in the case of the liberation movement from Western Sahara, a disputed territory partially annexed by Morocco in 1975. In exile in Algeria, Western Sahara's liberation movement has set up a state-like government that seeks international recognition as a state. In support of its efforts at state-making, the liberation movement has drawn on a longstanding local distinction within custom to produce a distinction between a‘rāf, construed as tribal laws to be erased, and ‘ādāt, construed as customary cultural heritage to be elevated