Habitat Selection and Movement of Pond-Breeding Amphibians in Experimentally Fragmented Pine Forests

ABSTRACT Population-level responses of amphibians to forest management regimes are partly dictated by individual behavioral responses to habitat alteration. We examined the short-term (i.e., 24-hr) habitat choices and movement patterns of 3 amphibian species—southern leopard frogs (Rana sphenocephala), marbled salamanders (Ambystoma opacum), and southern toads (Bufo terrestris)—released on edges between forest habitats and recent clear-cuts in the Upper Coastal Plain of South Carolina, USA. We predicted that adult frogs and salamanders would preferentially select forest using environmental cues as indicators of habitat suitability. We also predicted that movement patterns would differ in clear-cuts relative to forests, resulting in lower habitat permeability of clear-cuts for some or all of the species. Using fluorescent powder tracking, we determined that marbled salamanders selected habitat at random, southern toads preferred clear-cuts, and southern leopard frogs initially selected clear-cuts but ultimately preferred forests. Frogs exhibited long-distance, directional movement with few turns. In contrast, toads exhibited wandering behavior and salamanders moved relatively short distances before locating cover. Southern toads and southern leopard frogs moved farther in forests, and all 3 species made more turns in clear-cuts than in forests. Habitat selection by southern toads did not vary according to body size, sex, or the environmental cues we measured. However, marbled salamanders were more likely to enter clear-cuts when soil moisture was high, and southern leopard frogs were more likely to enter clear-cuts when relative humidity and air temperature were higher in the clear-cut than in adjacent forest. Although we found evidence of reduced habitat permeability of clear-cuts for southern leopard frogs and southern toads, none of the species exhibited strong behavioral avoidance of the small (4-ha) clear-cuts in our study. Further studies of long-term habitat use and the potential physiological and other costs to individuals in altered forests are needed to understand the effects of forest management on population persistence. To reduce potentially detrimental effects of clear-cutting on amphibians in the Southeast, wildlife managers should consider the vagility and behavior of species of concern, especially in relation to the size of planned harvests adjacent to breeding sites.     

Biodiversity and the Heterogeneous Disturbance Regime on Military Training Lands

Disproportionately large numbers of threatened and endangered species and unusually high biodiversity occur on active and former military training areas. Although this may seem paradoxical given the apparently destructive nature of military training, an evaluation of the nature and extent of the disturbances is enlightening. Military training frequently produces heterogeneous landscapes. Large portions of military training areas remain virtually untouched, favoring disturbance‐averse species; other portions are heavily disturbed, favoring disturbance‐dependent species. The rich habitat mosaics include the two extremes as well as the continua of disturbance and succession between them, thus providing suitable habitat for a very large number of species with widely varying habitat requirements. To explain the phenomenon, a heterogeneous disturbance hypothesis is proposed which suggests that biodiversity is maximized where multiple kinds, frequencies, severities, periodicities, sizes, shapes, and/or durations of disturbance occur concomitantly on a landscape in a spatially and temporally distributed fashion. The enhanced biodiversity occurring on active and former military training areas illustrates the need for restoration ecologists to restore or maintain an appropriate heterogeneous disturbance regime when attempting to restore ecosystem function and biodiversity.     

Habitat split as a driver of disease in amphibians

Anthropogenic habitat disturbance is fundamentally altering patterns of disease transmission and immunity across the vertebrate tree of life. Most studies linking anthropogenic habitat change and disease focus on habitat loss and fragmentation, but these processes often lead to a third process that is equally important: habitat split. Defined as spatial separation between the multiple classes of natural habitat that many vertebrate species require to complete their life cycles, habitat split has been linked to population declines in vertebrates, e.g. amphibians breeding in lowland aquatic habitats and overwintering in fragments of upland terrestrial vegetation. Here, we link habitat split to enhanced disease risk in amphibians (i) by reviewing the biotic and abiotic forces shaping elements of immunity and (ii) through a spatially oriented field study focused on tropical frogs. We propose a framework to investigate mechanisms by which habitat split influences disease risk in amphibians, focusing on three broad host factors linked to immunity: (i) composition of symbiotic microbial communities, (ii) immunogenetic variation, and (iii) stress hormone levels. Our review highlights the potential for habitat split to contribute to host-associated microbiome dysbiosis, reductions in immunogenetic repertoire, and chronic stress, that often facilitate pathogenic infections and disease in amphibians and other classes of vertebrates. We highlight that targeted habitat-restoration strategies aiming to connect multiple classes of natural habitats (e.g. terrestrial–freshwater, terrestrial–marine, marine–freshwater) could enhance priming of the vertebrate immune system through repeated low-load exposure to enzootic pathogens and reduced stress-induced immunosuppression.     

Active military training areas as refugia for disturbance-dependent endangered insects

Oedipoda caerulescens (blue-winged grasshopper) and Cicindela hybrida (northern dune tiger beetle) are protected insects in Germany and elsewhere. They are known to occur on sparsely vegetated, sandy soil. Populations of the two insects were evaluated in relation to physical soil disturbance on four military training areas in Germany to determine if the military disturbance regime occurring there is conducive to the survival of the species and to provide insight into the nature of the disturbance that may be necessary as conservationists seek ways to maintain, establish or re-establish suitable habitat. Adults of O. caerulescens exhibited statistically significant preference for areas with between 60% and 100% surface disturbance, corresponding to 50–70% plant cover, depending on the location. C. hybrida adults preferentially occupied areas with >40% disturbance resulting in an average of 61% plant cover. The results confirm suggestions that both species are disturbance-dependent. Military training areas represent some of the last, large remnants of sparse, dry, sandy grasslands in Europe. The nature of land-based military training creates suitable habitat patches as well as habitat connectivity needed for the maintenance of metapopulations. As a result, military training areas represent some of the last remaining vestiges of a habitat and disturbance regime that are highly favored by O. caerulescens and C. hybrida and other species with similar habitat requirements.     

Was macht Molche giftig?

What makes newts toxic? Tetrodotoxin is one of the strongest toxins of natural origin, which specifically blocks voltage-gated sodium channels of nerve and muscle. It was first detected in marine puffer fish (Tetraodontidae), but later also in other marine and terrestrial animals such as in amphibians. Particularly, some newt species (Urodela) of the New and Old World exhibit extremely high concentrations of the toxin which, to some extent, provide protection against predators. However, North American garter snakes of the genus Thamnophis proved to be toxin-resistant which is based on amino acid substitutions at the toxin-binding site of the sodium channel. The origin of the toxin is under discussion. Apparently, it is not synthesized by the newts but is most likely acquired via the food-chain and stored. Moreover, other amphibians such as some toads and frogs have been found to contain tetrodotoxin. The significance of the toxin for survival of species and its interweaving with the challenges of the ecosystems is still unclear.     

Sex and age differences in habitat use by invasive cane toads (Rhinella marina) and a native anuran (Cyclorana australis) in the Australian wet–dry tropics

Although generalized habitat use may contribute to the success of invasive taxa, even species that are typically described as habitat generalists exhibit non‐random patterns of habitat use. We measured abiotic and biotic factors in 42 plots (each 100 × 10 m) along a 4.2‐km long unpaved road in tropical Australia, at a site that had been invaded by cane toads (Rhinella marina Bufonidae) seven years previously. We also counted anurans at night in each of these plots on 103 nights during the tropical wet season, over a five‐year period, beginning soon after the initial toad invasion. Spatial distributions differed significantly among adult male toads (n = 1047), adult female toads (n = 1222), juvenile toads (n = 342) and native frogs (Cyclorana australis Hylidae, n = 234). Adult male toads were closely associated with water bodies used as calling and/or spawning sites, whereas adult female toads and native frogs were most commonly encountered in drier forested areas on sloping ground. Juvenile toads used the margins of the floodplain more than conspecific adults did, but the floodplain itself was rarely used. Understanding which components of the habitat are most important to specific age and sex classes within a population, or how invasive species differ from native species in this respect, can clarify issues such as the spatial and temporal location of ecological impact by an invader, and the most effective places for control of the invader with minimal collateral effects on the native biota.     

Effects of an invasive species on refuge‐site selection by native fauna: The impact of cane toads on native frogs in the Australian tropics

Invasive species can induce shifts in habitat use by native taxa: either by modifying habitat availability, or by repelling or attracting native species to the vicinity of the invader. The ongoing invasion of cane toads (Rhinella marina) through tropical Australia might affect native frogs by affecting refuge‐site availability, because both frogs and toads frequently shelter by day in burrows. Our laboratory and field studies in the wet‐dry tropics show that native frogs of at least three species (Litoria tornieri, Litoria nasuta and Litoria dahlii) preferentially aggregate with conspecifics, and with (some) other species of native frogs. However, the frogs rarely aggregated with cane toads either in outdoor arenas or in standardized experimental burrows that we monitored in the field. The native frogs that we tested either avoided burrows containing cane toads (or cane toad scent) or else ignored the stimulus (i.e. treated such a burrow in the same way as they did an empty burrow). Native frogs selected a highly non‐random suite of burrows as diurnal retreat sites, whereas cane toads were less selective. Hence, even in the absence of toads, frogs do not use many of the burrows that are suitable for toads. The invasion of cane toads through tropical Australia is unlikely to have had a major impact on retreat‐site availability for native frogs.     

Species‐specific responses to wetland mitigation among amphibians in the Greater Yellowstone Ecosystem

Habitat loss and degradation are leading causes of biodiversity declines, therefore assessing the capacity of created mitigation wetlands to replace habitat for wildlife has become a management priority. We used single season occupancy models to compare the occurrence of larvae of four species of pond‐breeding amphibians in wetlands created for mitigation, wetlands impacted by road construction, and unimpacted reference wetlands along a highway corridor in the Greater Yellowstone Ecosystem, United States. Created wetlands were shallow and had less aquatic vegetation and surface area than impacted and reference wetlands. Occupancy of barred tiger salamander (Ambystoma mavortium) and boreal chorus frog (Pseudacris maculata) larvae was similar across wetland types, whereas boreal toads (Anaxyrus boreas) occurred more often in created wetlands than reference and impacted wetlands. However, the majority of created wetlands (>80%) dried partially or completely before amphibian metamorphosis occurred in both years of our study, resulting in heavy mortality of larvae and, we suspect, little to no recruitment. Columbia spotted frogs (Rana luteiventris), which require emergent vegetation that is not common in newly created wetlands, occurred commonly in impacted and reference wetlands but were found in only one created wetland. Our results show that shallow created wetlands with little aquatic vegetation may be attractive breeding areas for some amphibians, but may result in high mortality and little recruitment if they fail to hold water for the entire larval period.     

The Last South American Redoubt? Tested Surinamese Anurans Still Chytrid Free

Batrachochytrium dendrobatidis, a chytrid fungus infecting amphibians’ cutaneous layer, is responsible for the greatest contemporary loss of amphibian biodiversity. In South America, Suriname is one of the only three countries where B. dendrobatidis infections of anurans (frogs and toads) have not been documented. To further examine this apparent gap in pathogen occurrence, frogs were sampled for B. dendrobatidis spores at eight disparate geographic locations in Suriname, including locations with high and low levels of anthropogenic activities, and near Suriname’s border with Brazil and French Guiana, countries where B. dendrobatidis infections have been documented. None of the 347 frogs sampled, representing 37 species from eight families, tested positive for B. dendrobatidis. Our results provide the baseline data for future comparative testing and one of the last opportunities for a country in South America to proactively plan mitigation measures to protect amphibians from B. dendrobatidis’ presumed eventual incursion into Suriname.

     

Spawning site selection by feral cane toads (Bufo marinus) at an invasion front in tropical Australia

Abstract Spawning sites are a critical and often scarce resource for aquatic‐breeding amphibians, including invasive species such as the cane toad (Bufo marinus). If toads select spawning sites based on habitat characteristics, we can potentially manipulate those characteristics to either enhance or reduce their suitability as breeding sites. We surveyed 25 spawning sites used by cane toads, and 25 adjacent unused sites, in an area of tropical Australia recently invaded by these feral anurans. Water chemistry (pH, conductivity, salinity, turbidity) was virtually identical between the two sets of waterbodies, but habitat characteristics were very different. Toads selectively oviposited in shallow pools with gradual rather than steep slopes, and with open (unvegetated) gradually sloping muddy banks. They avoided flowing water, and pools with steep surrounds. In these respects, cane toads broadly resemble previously studied toad species in other parts of the world, as well as conspecifics within their natural range in South America.     

Multi-event models reveal the absence of interaction between an invasive frog and a native endangered amphibian

Among the multiple factors involved in the decline of amphibians, competition with alien species remains understudied. In Western Europe, non-native marsh frogs (Pelophylax ridibundus and other sister species) have been widely imported for gastronomic purposes and have then often escaped from captivity. Apart from closely related species, the impact of the subsequent colonization of non-native species on native amphibians is still unknown. In this study, we analysed the response of a threatened species, the yellow-bellied toad (Bombina variegata), faced with the presence of alien frogs in mountain rivers in France. We studied the co-occurrence pattern of the native toad and the alien frog, while taking into account co-dependence of the detection rates in both species. We tested three main scenarios to explain a non-random pond occurrence for each species and to establish their respective habitat preferenda: (1) a pond-area-dependent scenario, predicting different responses from each species for a given pond size, (2) a fish-dependent scenario, predicting different responses from each species given the presence of fish in ponds, and (3) a floodplain-width-dependent scenario, predicting different responses from each species given the geomorphological characteristics of the floodplain. Taking into account site-specific covariates, we concluded that introduced frogs do not currently have an impact on the native population of the yellow-bellied toad.     

Density dependence in the terrestrial life history stage of two anurans

Populations of species with complex life cycles have the potential to be regulated at multiple life history stages. However, research tends to focus on single stage density-dependence, which can lead to inaccurate conclusions about population regulation and subsequently hinder conservation efforts. In amphibians, many studies have demonstrated strong effects of larval density and have often assumed that populations are regulated at this life history stage. However, studies examining density regulation in the terrestrial stages are rare, and the functional relationships between terrestrial density and vital rates in amphibians are unknown. We determined the effects of population density on survival, growth and reproductive development in the terrestrial stage of two amphibians by raising juvenile wood frogs (Rana sylvatica) and American toads (Bufo americanus) at six densities in terrestrial enclosures. Density had strong negative effects on survival, growth and reproductive development in both species. We fitted a priori recruitment functions to describe the relationship between initial density and the density of survivors after one year, and determined the functional relationship between initial density and mass after one year. Animals raised at the lowest densities experienced growth and survival rates that were over twice as great as those raised at the highest density. All female wood frogs in the lowest density treatment showed signs of reproductive development, compared to only 6% in the highest density treatment. Female American toads reached minimum reproductive size only at low densities, and male wood frogs and American toads reached maturity only in the three lowest density treatments. Our results demonstrate that in the complex life cycle of amphibians, density in the terrestrial stage can reduce growth, survival and reproductive development and may play an important role in amphibian population regulation. We discuss the implications of these results for population regulation in complex life cycles and for amphibian conservation.     

Amphibian road mortality in Europe: a meta-analysis with new data from Poland

While the increasing vehicular traffic is widely suspected to play a role in the worldwide amphibian population decline, the research of amphibian road mortality is scarce, fragmented, fraught with methodological problems, and largely inconclusive. As the first attempt at a synthesis, we analyzed all available data on amphibian mortality in Europe and combined them with four previously unpublished surveys conducted by us. Based on our recalculation of road-kill counts in terms of species-specific road-kill recordability, we conclude that, in lowland Central Europe, the common toads, Bufo bufo, are the most common victims of vehicular traffic in suburban landscapes, while the common frogs, Rana temporaria and Triturus newts, prevail in rural landscapes. The green frogs also tend to be more frequent in rural areas. Common tree frogs, Hyla arborea, are unexpectedly rare in the road-kill record despite their terrestrial and migratory habits. In consideration of problems with obtaining accurate amphibian population estimates, we further propose the road kills-to-spawners ratio (R/S) as a working measure of the impact of road mortality on a local population. While the R/S ratio may not reflect the losses to an entire local amphibian population, it is free of the errors of whole-population estimates, which are notoriously difficult for amphibians. When corrected for species-specific road-kill recordability, most results suggest that the impact of roads on newts may have been underestimated and that the impact on common frog populations may be higher than on those of common toads. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Communicated by H. Kierdorf     

Chytridiomycosis Widespread in Anurans of Northeastern United States

ABSTRACT An emerging disease of amphibians caused by the chytrid fungus Batrachochytrium dendrobatidis has been associated with morbidity, mortality, and extinction of species. Typically, researchers have detected B. dendrobatidis only when examining amphibians for causes of mortalities; few data exist on infection rates where mortalities are lacking. During May-September 2000–2002 we obtained amphibian specimens killed by vehicles and others collected at remote off-road sites throughout Maine, USA, and from federal lands in 5 states in the Northeast. We detected infected specimens, mostly green frogs (Rana clamitans), at 5 of 7 national wildlife refuges, a federal waterfowl production area, and Acadia National Park. Seven of 9 species, including all Ranidae species, were infected throughout Maine; rates ranged from 14.6% in American toads (Bufo americanus) to 25.7% in northern leopard frogs (Rana pipiens). We did not detect any infections in 50 eastern gray tree frogs (Hyla versicolor) or 21 spring peepers (Pseudacris crucifer). Species that hibernate in terrestrial habitats seem to have lower rates of infection than species that hibernate in aquatic habitats. Infections peaked in spring and autumn and were associated with air temperatures optimal for B. dendrobatidis growth. The relatively high infection rates among species without documented die-offs suggest that either losses have occurred undetected, that the fungus is endemic and species have attained a level of resistance to infections becoming lethal, or that climatic conditions of the Northeast have a role in preventing infections from being lethal. Data on prevalence and distribution of this chytrid fungus in the Northeast may be useful in modeling its origins and predicting long-term ecosystem effects involving anurans.     

Identifying core habitat before it’s too late: the case of Bombina variegata,an internationally endangered amphibian

Impending land-use change, including agricultural intensification, is increasingly threatening biodiversity in traditional rural landscapes. To ensure the persistence of species that are vulnerable to land-use change it is necessary to identify and protect high quality habitat before species start to decline. Given that many potentially vulnerable species are still widespread in traditional rural landscapes, it is difficult to identify particularly important locations for such species. Presence–absence data on a given species may have limited application in such cases. As an alternative to presence–absence data, we investigated the influence of environmental variables on the physiological body condition of Bombina variegata (yellow-bellied toad) in a traditional rural landscape in Transylvania, Romania. The species is internationally endangered but remains common throughout our study area. Based on body condition measurements of 550 toads from 60 ponds, we found that toads in forest ponds had significantly better body condition than those in pasture ponds, indicating that forest landscapes provided particularly high quality habitat. We suggest that measures such as body condition—in addition to distribution data—could have considerable application in identifying high quality habitat for other species that are still widespread in traditional landscapes, but have declined in modernised, but otherwise similar landscapes.     

Behavioural tactics used by invasive cane toads (Rhinella marina) to exploit apiaries in Australia

Behavioural flexibility plays a key role in facilitating the ability of invasive species to exploit anthropogenically‐created resources. In Australia, invasive cane toads (Rhinella marina) often gather around commercial beehives (apiaries), whereas native frogs do not. To document how toads use this resource, we spool‐tracked cane toads in areas containing beehives and in adjacent natural habitat without beehives, conducted standardized observations of toad feeding behaviour, and ran prey‐manipulation trials to compare the responses of cane toads versus native frogs to honeybees as potential prey. Toads feeding around beehives travelled shorter distances per night, and hence used different microhabitats, than did toads from nearby control sites without beehives. The toads consumed live bees from the hive entrance (rather than dead bees from the ground), often climbing on top of one another to gain access to the hive entrance. Prey manipulation trials confirm that bee movement is the critical stimulus that elicits the toads’ feeding response; and in standardized trials, native frogs consumed bees less frequently than did toads. In summary, cane toads flexibly modify their movements, foraging behaviour and dietary composition to exploit the nutritional opportunities created by commercial beehives, whereas native anurans do not.     

Native Australian frogs avoid the scent of invasive cane toads

Invasive species can affect the ecosystems they colonize by modifying the behaviour of native taxa; for example, avoidance of chemical cues from the invader may modify habitat use (shelter site selection) by native species. In laboratory trials, we show that metamorphs of most (but not all) native frog species on a tropical Australian floodplain avoid the scent of invasive cane toads (Bufo marinus Linnaeus 1758). Cane toads also avoid conspecific scent. This response might reduce vulnerability of metamorph frogs and toads to larger predatory toads. However, similar avoidance of one type of pungency control (garlic), and the presence of this avoidance behaviour in frogs at the toad invasion front (and hence, with no prior exposure to toads), suggest that this may not be an evolved toad‐specific response. Instead, our data support the simpler hypothesis that the metamorph anurans tend to avoid shelter sites that contain strong and unfamiliar scents. Temporal and spatial differences in activity of frogs versus toads, plus the abundance of suitable retreat sites during the wet season (the primary time of frog activity), suggest that avoiding toad scent will have only a minor impact on the behaviour of native frogs. However, this behavioural impact may be important when environmental conditions bring toads and frogs into closer contact.     

RESTRICTED GENE FLOW IN A MOVING HYBRID ZONE OF THE NEWTS TRITURUS CRISTATUS AND T. MARMORATUS IN WESTERN FRANCE

Two hybridizing species of newts, Triturus cristatus and T. marmoratus, with overlapping distributions show a parapatric distribution when surveyed in detail. The factors that govern the distribution of cristatus vs. marmoratus in the département (province) of Mayenne in western France are identified as forestation and relief. The parapatric hybrid zone running through Mayenne is narrow but widens to approximately 20 km in an area with mixed habitat. In this area most breeding sites are shared and F₁ hybrids form about 4% of the total population. Analysis of survey data collected about 30 years previously also shows an essentially parapatric distribution. Comparison of past and present distribution maps reveals that cristatus has superseded marmoratus over large areas in the south of Mayenne. An area where marmoratus replaced cristatus also exists, but it is more limited in size. Gene flow between cristatus and marmoratus is analyzed using 10 diagnostic genetic markers [9 protein loci and mitochondrial (mt) DNA]. In syntopic populations nuclear gene flow is bidirectional with a mean frequency of introgressed alleles (f) of 0.3%. In allotopic populations of cristatus and marmoratus gene flow is present in areas of species replacement (f = 0.3%), while gene flow appears to be absent in those areas that have been continuously occupied by a single species. At the biogeographic level, the presence or absence of introgression is paralleled by the persistence or absence, respectively, of pockets of cristatus–marmoratus syntopy. All F₁ hybrids possess the cristatus type mtDNA. This may be due to asymmetric interspecific mate choice and would explain the observed absence of introgression of the maternally inherited mtDNA genome in areas where cristatus replaced marmoratus. The cristatus–marmoratus hybrid zone bears characteristics of both the clinal (parapatric) hybrid zone model and the mosaic hybrid zone model. Such a mixed model—for which we propose the term “reticulate hybrid zone”—can be appreciated only if studied over a two-dimensional geographic area and also through time.     

Global amphibian declines: a perspective from the Caribbean

Recent concern over the possibility of a global decline in amphibians prompted this assessment of the West Indian species. At the species level, the West Indian amphibian fauna (156 species, all frogs and toads) has not undergone a general decline, and no species is known to be extinct. However, one Puerto Rican species (Eleutherodactylus karlschmidti) has not been seen in over ten years despite considerable search effort. Seven other species, including the Puerto Rican livebearing frog (E. jasperi), have not been seen recently, although their present status cannot be determined until additional effort is made to locate them. Two stream-associated species on Hispaniola (E. semipalmatus and Hyla vasta) appear to have declined in recont years, probably due to the alteration of riparian habitats by deforestation. Other vertebrate groups in the West Indies, such as mammals, have been more affected by human-caused environmental degradation than have amphibians. Large-scale extinctions of frogs and other forest-dwelling species are not expected to occur until forest cover reaches very low levels. Haiti is on the brink of such extinctions with less than 1% of its forest cover remaining. Two recommendations are made to help curtail the expected loss of biodiversity: (i) import charcoal to replace that produced by burning native trees (used as cooking fuel), as an immediate measure, and (ii) control human population growth, as a long-term solution.     

Quantifying Anuran Microhabitat Use to Infer the Potential for Parasite Transmission between Invasive Cane Toads and Two Species of Australian Native Frogs

Parasites that are carried by invasive species can infect native taxa, with devastating consequences. In Australia, invading cane toads (Rhinella marina) carry lungworm parasites (Rhabdias pseudosphaerocephala) that (based on previous laboratory studies) can infect native treefrogs (Litoria caerulea and L. splendida). To assess the potential of parasite transmission from the invader to the native species (and from one infected native frog to another), we used surveys and radiotelemetry to quantify anuran microhabitat use, and proximity to other anurans, in two sites in tropical Australia. Unsurprisingly, treefrogs spent much of their time off the ground (especially by day, and in undisturbed forests) but terrestrial activity was common at night (especially in anthropogenically modified habitats). Microhabitat overlap between cane toads and frogs was generally low, except at night in disturbed areas, whereas overlap between the two frog species was high. The situations of highest overlap, and hence with the greatest danger of parasite transmission, involve aggregations of frogs within crevices by day, and use of open ground by all three anuran species at night. Overall, microhabitat divergence between toads and frogs should reduce, but not eliminate, the transmission of lungworms from invasive toads to vulnerable native frogs.