Invasive Toads Research Articles

Constructing an Invasion Machine: The Rapid Evolution of a Dispersal-Enhancing Phenotype During the Cane Toad Invasion of Australia.

Biological invasions can induce rapid evolutionary change. As cane toads (Rhinella marina) have spread across tropical Australia over an 80-year period, their rate of invasion has increased from around 15 to 60 km per annum. Toads at the invasion front disperse much faster and further than conspecifics from range-core areas, and their offspring inherit that rapid dispersal rate. We investigated morphological changes that have accompanied this dramatic acceleration, by conducting three-dimensional morphometric analyses of toads from both range-core and invasion-front populations. Morphology of heads, limbs, pectoral girdles and pelvic girdles differed significantly between toads from the two areas, ranging from 0.5% to 16.5% difference in mean bone dimensions between populations, with invasion-front toads exhibiting wider forelimbs, narrower hindlimbs and more compact skulls. Those changes plausibly reflect an increased reliance on bounding (multiple short hops in quick succession) rather than separate large leaps. Within an 80-year period, invasive cane toads have converted the basic anuran body plan – which evolved for occasional large leaps to evade predators – into a morphotype better-suited to sustained long-distance travel.

Using genetic data to predict the vulnerability of a native predator to a toxic invader

Australia has no native toad species, and as a consequence, many Australian pre - dators lack resistance to the toxins of the invasive cane toad Rhinella marina, and die if they ingest one of these toads. Resistance is conferred by a small and consistent genetic change, so genetic data can provide a rapid, non-invasive way to clarify the vulnerability of as-yet-unstudied taxa. To evaluate the hypothesis that a recent decline of ghost bat Macroderma gigas populations in tropical Australia is due to ingestion of cane toads, we sequenced the H1-H2 extracellular domain of the sodium-potassium-ATPase. Two anuran-eating Asian relatives of the Australian species possess the genes that confer bufotoxin resistance, but the ghost bat does not. Like varanid lizards (major victims of the toad invasion), Australian ghost bats appear to have lost their physiological resistance to toad toxins but retained generalist foraging behaviours, potentially including a readiness to attack toads as well as frogs. Our genetic data suggest that cane toads may imperil populations of this iconic predator, and detailed behavioural and ecological studies are warranted.

Dietary overlap between the invasive coquí frog (Eleutherodactylus coqui) and the Hawaiian hoary bat (Lasiurus cinereus semotus) on the Island of Hawai’i

The invasive coqui frog is a likely insectivorous competitor to the native Hawaiian hoary bat. The frog is a sit-and-wait predator native to Puerto Rico, but it has the capacity for producing dense populations in its invasive range and the potential to reduce arthropod populations, including aerial arthropods in the orders Coleoptera, Lepidoptera and Isoptera, which are primary food sources of the Hawaiian hoary bat. Dietary analysis of the coqui frog showed that aerial insects made up 33.8 % of the diet. The dietary similarity of the bat and frog was relatively low (Pianka index = 0.25), however, coqui frogs consumed a wide range of juvenile flying insects. Population densities of coqui frogs at two low elevation sites were projected to range from 750 to 14,000 individuals/hectare, and could consume an estimated 1500–19,000 aerial insects/hectare/night, or nearly 90.6 % of all available aerial insects/hectare. Although direct competition between coqui frogs and Hawaiian hoary bats was not confirmed, the coqui frogs could reduce available prey for bats in newly invaded upland sites where bats appear to be more selective of prey species among available aerial insects.

Rapid differentiation of sexual signals in invasive toads: call variation among populations.

Advertisement calls tend to differ among populations, based on morphological and environmental factors, or simply geographic distance, in many taxa. Invasive cane toads (Rhinella marina) were introduced to Australia in 1935 and their distribution has expanded at increasing rates over time. Rapid evolution occurred in morphological and behavioural characters that accelerate dispersal, but the effects of rapid expansion on sexual signals have not been examined. We collected advertisement calls from four populations of different ages since invasion, and analysed the geographic differentiation of seven call parameters. Our comparisons indicate that the calls of R. marina differ among Australian populations. The signal variation was not simply clinal with respect to population age, climate, or morphological differentiation. We suggest that selection on signalling among populations has been idiosyncratic and may reflect local female preferences or adaptation to environmental factors that are not clinal such as energy availability.

Frogs in the spotlight: a 16-year survey of native frogs and invasive toads on a floodplain in tropical Australia.

Although widespread declines in anuran populations have attracted considerable concern, the stochastic demographics of these animals make it difficult to detect consistent trends against a background of spatial and temporal variation. To identify long‐term trends, we need datasets gathered over long time periods, especially from tropical areas where anuran biodiversity is highest. We conducted road surveys of four anurans in the Australian wet–dry tropics on 4637 nights over a 16‐year period. Our surveys spanned the arrival of invasive cane toads (Rhinella marina), allowing us to assess the invader's impact on native anuran populations. Our counts demonstrate abrupt and asynchronous shifts in abundance and species composition from one year to the next, not clearly linked to rainfall patterns. Typically, periods of decline in numbers of a species were limited to 1–2 years and were followed by 1‐ to 2‐year periods of increase. No taxa showed consistent declines over time, although trajectories for some species showed significant perturbations coincident with the arrival of toads. None of the four focal frog species was less common at the end of the study than at the beginning, and three of the species reached peak abundances after toad arrival. Survey counts of cane toads increased rapidly during the initial stage of invasion but have subsequently declined and fluctuated. Distinguishing consistent declines versus stochastic fluctuations in anuran populations requires extensive time‐series analysis, coupled with an understanding of the shifts expected under local climatic conditions. This is especially pertinent when assessing impacts of specific perturbations such as invasive species.

Food Habits of Invasive Bullfrogs and Native Thin-Toed Frogs Occurring in Sympatry in Southeastern Brazil

Abstract. Although the feeding ecology of sympatric anurans has been studied, less is known about the food habits of co-occurring invasive and native frog species, particularly in the Neotropics. We compared the food habits of invasive bullfrogs (Lithobates catesbeianus) and native thin-toed frogs (Leptodactylus cf. latrans) in a locality of Southeastern Brazil. Monthly field surveys and stomach content analyses were performed between September 2008–April 2009 and September–November 2009. Prey items were identified to the lowest taxonomic level and diet overlap was estimated. Juveniles of both species had similar body sizes, but adult thin-toed frogs were smaller than adult bullfrogs. Insects and spiders dominated the diets of juveniles of both species and of adult thin-toed frogs, whereas anurans and water bugs predominated in adult bullfrog diets. Thin-toed frogs consumed a larger number of prey items than bullfrogs, which in turn consumed more voluminous prey items. Aquatic and amphibious prey items we...

Can we control the invasive cane toad using chemicals that have evolved under intraspecific competition?

Many invasive species experience intense intraspecific competition, because they are abundant in anthropogenically disturbed habitats where few native species persist. Species-specific competitive mechanisms that evolve in this context may offer novel, highly targeted means to control invasive taxa. We conducted laboratory experiments to evaluate the feasibility of this method of control, based on waterborne cues that are produced by tadpoles of the cane toad (Rhinella marina) to suppress the development of conspecific embryos. Our trials examined the nature and species-specificity of the effect, the robustness of the cue to freezing and storage, and the amounts required to suppress toad embryos. Our results were encouraging. The cue appears to be chemical rather than a biological organism, and may well be species-specific; the four species of native anurans that we tested were not influenced by toad larval cues. The cue retains its effectiveness after being frozen, but not after being dried, or after 7 d in water. It is effective at very low concentrations (the amount produced by three tadpoles within 750 L of water). Overall, the cane toad's suppressor pheromone may offer an effective new way to control invasive toads.

Geographical variations in adult body size and reproductive life history traits in an invasive anuran, Discoglossus pictus

Variability in life history traits positively affects the establishment and expansive potential of invasive species. In the present study, we analysed the variation of body size in seven populations − two native and five invasive − of the painted frog (Discoglossus pictus, Anura: Discoglossidae), native to North Africa and introduced in southern France and the north-east of the Iberian Peninsula. Other life history traits (age at maturity, size at maturity, longevity, median age and potential reproductive lifespan) were analysed in a native and an invasive population. We observed geographic variations in adult body size, related mainly to mean annual precipitation. Thus, populations had greater body size as mean annual precipitation increased, resulting in bigger specimens in the invasive populations. Adult body size and growth rates also varied between sexes in all studied populations, with males significantly larger than females. Age distribution varied between native (1–5 years) and invasive populations (2–4 years) and also between sexes. Our results suggest that higher precipitation promotes faster growth rates and larger adult body size that could facilitate the successful establishment of invasive populations.

Functional distance to recipient communities may favour invasiveness: insights from two invasive frogs

AbstractAimInvasive species present negative impacts on native biodiversity at a global scale. A key goal of community ecology is to identify what drives invasiveness, but hypotheses relying on biotic mechanisms remain largely untested for many groups. Here we asked whether source and recipient communities of two highly successful invasive anurans (the bullfrog Lithobates catesbeianus and the cane toad Rhinellla marina) differ consistently from a taxonomic and/or functional standpoint. If affirmative, this pattern could suggest that taxonomic and/or functional distances between an invasive species and a potentially recipient community might influence the alien's invasive potential.LocationWorld‐wide.MethodsBased on co‐occurrence data of 1061 amphibian species, we compared 30 source to 30 recipient communities of bullfrogs and cane toads by means of biotic metrics that summarize taxonomic and functional diversity and the relative position of the invasive species within the community. We also included environmental drivers that reportedly influence invasibility (climate, resource availability, spatial heterogeneity, and propagule pressure).ResultsBoth invasive species were functionally distant to their respective recipient communities; in contrast, community diversity did not explain much variation between source and recipient communities. Climate matching possibly influenced cane toad's but not bullfrog's invasiveness, and landscape factors had little relevance overall.Main conclusionThis study advances the notion that the relative position of a recently introduced species within the native functional space may help predicting its invasive potential.

Ecological immunization: in situ training of free-ranging predatory lizards reduces their vulnerability to invasive toxic prey.

In Australia, large native predators are fatally poisoned when they ingest invasive cane toads (Rhinella marina). As a result, the spread of cane toads has caused catastrophic population declines in these predators. Immediately prior to the arrival of toads at a floodplain in the Kimberley region, we induced conditioned taste aversion in free-ranging varanid lizards (Varanus panoptes), by offering them small cane toads. By the end of the 18-month study, only one of 31 untrained lizards had survived longer than 110 days, compared to more than half (nine of 16) of trained lizards; the maximum known survival of a trained lizard in the presence of toads was 482 days. In situ aversion training (releasing small toads in advance of the main invasion front) offers a logistically simple and feasible way to buffer the impact of invasive toads on apex predators.

Thermal ecological physiology of native and invasive frog species: do invaders perform better?

Biological invasions are recognized as an important biotic component of global change that threatens the composition, structure and functioning of ecosystems, resulting in loss of biodiversity and displacement of native species. Although ecological characteristics facilitating the establishment and spread of non-native species are widely recognized, little is known about organismal attributes underlying invasion success. In this study, we tested the effect of thermal acclimation on thermal tolerance and locomotor performance in the invasive Xenopus laevis and the Chilean native Calyptocephalella gayi. In particular, the maximal righting performance (μMAX), optimal temperature (TO), lower (CTmin) and upper critical thermal limits (CTmax), thermal breadth (Tbr) and the area under the performance curve (AUC) were studied after 6weeks acclimation to 10 and 20°C. We observed higher values of μmax and AUC in X. laevis in comparison to C. gayi. On the contrary, the invasive species showed lower values of CTmin in comparison to the native one. In contrast, CTmax, TO and Tbr showed no inter-specific differences. Moreover, we found that both species have the ability to acclimate their locomotor performance and lower thermal tolerance limit at low temperatures. Our results demonstrate that X. laevis is a better performer than C. gayi. Although there were differences in CTmin, the invasive and native frogs did not differ in their thermal tolerance. Interestingly, in both species the lower and upper critical thermal limits are beyond the minimal and maximal temperatures encountered in nature during the coldest and hottest month, respectively. Overall, our findings suggest that both X. laevis and C. gayi would be resilient to climate warming expectations in Chile.

Living up to its name? The effect of salinity on development, growth, and phenotype of the "marine" toad (Rhinella marina).

The highly permeable integument of amphibians renders them vulnerable to chemical characteristics of their environment, especially during the aquatic larval stage. As the cane toad (Rhinella marina, Bufonidae) invades southwards along the east coast of Australia, it is encountering waterbodies with highly variable conditions of temperature, pH, and salinity. Understanding the tolerance of toads to these conditions can clarify the likely further spread of the invader, as well as the adaptability of the species to novel environmental challenges. We measured salinity in waterbodies in the field and conducted laboratory trials to investigate the impacts of salinity on toad viability. Eggs and tadpoles from the southern invasion front tolerated the most saline conditions we found in potential spawning ponds during surveys [equivalent to 1200 ppm (3.5 % the salinity of seawater)]. Indeed, high-salinity treatments increased tadpole body sizes, accelerated metamorphosis, and improved locomotor ability of metamorphs (but did not affect metamorph morphology). At very low salinity [40 ppm (0.1 % seawater)], eggs hatched but larvae did not develop past Gosner stage 37. Our study shows that the egg and larval life stages of cane toads can tolerate wide variation in the salinity of natal ponds and that this aspect of waterbody chemistry is likely to facilitate rather than constrain continued southward expansion of the toad invasion front in eastern Australia.

Comparative dietary ecology of turtles (Chelodina burrungandjii and Emydura victoriae) across the Kimberley Plateau, Western Australia, prior to the arrival of cane toads

Food webs in north-western Australian rivers exist in dynamic environments and will be influenced by land use practices, invasion of toxic cane toads (Rhinella marina) and the effects of climate change on river flows. Baseline studies are needed to understand aquatic food webs before these impacts. In the present study, we investigated the diets of two turtles (Emydura victoriae and Chelodina burrungandjii) in four upland rivers across a gradient of rainfall and land uses in the Kimberley Plateau of Western Australia. We captured turtles by snorkelling and recovered their prey by stomach lavage. We enumerated 2720 prey items from 390 E. victoriae samples and 308 prey items from 155 C. burrungandjii samples. Prey compositions distinguished E. victoriae as an omnivorous generalist relying on a diversity of animal and plant prey and C. burrungandjii as a piscivorous specialist, but with both species as likely predators of toxic cane toad eggs or tadpoles. Comparisons among the rivers showed variation in diets for both species that reflect differences in prey availability and location-specific food webs. Terrestrially based food sources were observed in 26% of E. victoriae samples and 3% of C. burrungandjii samples, which indicates the importance of the aquatic–terrestrial interface and land use practices within these rivers.

The impact of lungworm parasites on rates of dispersal of their anuran host, the invasive cane toad

Translocation of native-range parasites to control invasive species is effective only if the parasite substantially impairs either the viability or dispersal rate of the invasive host. Lungworms (Rhabdias pseudosphaerocephala) of cane toads (Rhinella marina) were introduced to Australia from the toad’s native range, along with the toads, and have been suggested as a potential biocontrol of invasive toads due to various negative impacts on toad viability. We conducted two radio-telemetry studies on a tropical floodplain to specifically assess the parasite’s impact on toad dispersal. First, a retrospective correlative analysis of data from field-collected animals showed that toads infected with lungworms moved farther, not less, than uninfected conspecifics. Second, an experimental study (comparing movements of experimentally infected toads vs. uninfected controls) showed that lungworms did not modify rates of toad dispersal. In addition, experimental infection with lungworms did not elicit an immune response substantial enough to influence dispersal behaviour. Thus, we conclude that increasing lungworm densities at the invasion front as an attempt at biocontrol would not slow down the spread of cane toads.

Invasive toads shift predator-prey densities in animal communities by removing top predators.

Although invasive species can have substantial impacts on animal communities, cases of invasive species facilitating native species by removing their predators have rarely been demonstrated across vertebrate trophic linkages. The predictable spread of the invasive cane toad (Rhinella marina), however, offered a unique opportunity to quantify cascading effects. In northern Australia, three species of predatory monitor lizards suffered severe population declines due to toad-induced lethal toxic ingestion (yellow-spotted monitor (Varanus panoptes), Mertens' water monitor (V. mertensi), Mitchell's water monitor (V. mitchelli). We, thus, predicted subsequent increases in the abundance and recruitment of prey species due to the reduction of those predators. Toad-induced population-level declines in the water monitor species approached 50% over a five-year period spanning the toad invasion, apparently causing fledging success of the Crimson Finch (Neochmia.phaeton) to increase from 55% to 81%. The consensus of our original and published long-term data is that invasive cane toads are causing predators to lose a foothold on top-down regulation of their prey, triggering shifts in the relative densities of predator and prey in the Australian tropical savannah ecosystem.

Frog eat frog: exploring variables influencing anurophagy.

Background. Frogs are generalist predators of a wide range of typically small prey items. But descriptions of dietary items regularly include other anurans, such that frogs are considered to be among the most important of anuran predators. However, the only existing hypothesis for the inclusion of anurans in the diet of post-metamorphic frogs postulates that it happens more often in bigger frogs. Moreover, this hypothesis has yet to be tested.Methods. We reviewed the literature on frog diet in order to test the size hypothesis and determine whether there are other putative explanations for anurans in the diet of post-metamorphic frogs. In addition to size, we recorded the habitat, the number of other sympatric anuran species, and whether or not the population was invasive. We controlled for taxonomic bias by including the superfamily in our analysis.Results. Around one fifth of the 355 records included anurans as dietary items of populations studied, suggesting that frogs eating anurans is not unusual. Our data showed a clear taxonomic bias with ranids and pipids having a higher proportion of anuran prey than other superfamilies. Accounting for this taxonomic bias, we found that size in addition to being invasive, local anuran diversity, and habitat produced a model that best fitted our data. Large invasive frogs that live in forests with high anuran diversity are most likely to have a higher proportion of anurans in their diet.Conclusions. We confirm the validity of the size hypothesis for anurophagy, but show that there are additional significant variables. The circumstances under which frogs eat frogs are likely to be complex, but our data may help to alert conservationists to the possible dangers of invading frogs entering areas with threatened anuran species.

The impact of invasive cane toads on native wildlife in southern Australia.

Commonly, invaders have different impacts in different places. The spread of cane toads (Rhinella marina: Bufonidae) has been devastating for native fauna in tropical Australia, but the toads' impact remains unstudied in temperate-zone Australia. We surveyed habitat characteristics and fauna in campgrounds along the central eastern coast of Australia, in eight sites that have been colonized by cane toads and another eight that have not. The presence of cane toads was associated with lower faunal abundance and species richness, and a difference in species composition. Populations of three species of large lizards (land mullets Bellatorias major, eastern water dragons Intellagama lesueurii, and lace monitors Varanus varius) and a snake (red-bellied blacksnake Pseudechis porphyriacus) were lower (by 84 to 100%) in areas with toads. The scarcity of scavenging lace monitors in toad-invaded areas translated into a 52% decrease in rates of carrion removal (based on camera traps at bait stations) and an increase (by 61%) in numbers of brush turkeys (Alectura lathami). The invasion of cane toads through temperate-zone Australia appears to have reduced populations of at least four anurophagous predators, facilitated other taxa, and decreased rates of scavenging. Our data identify a paradox: The impacts of cane toads are at least as devastating in southern Australia as in the tropics, yet we know far more about toad invasion in the sparsely populated wilderness areas of tropical Australia than in the densely populated southeastern seaboard.

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.

Long‐term monitoring reveals declines in an endemic predator following invasion by an exotic prey species

AbstractInvasive predators can cause population declines in native prey species, but empirical evidence linking declines of native predators to invasive prey is relatively rare. Here, we document declines in an Australian freshwater crocodile Crocodylus johnstoni population following invasion of a toxic prey species, the cane toad Rhinella marina. Thirty‐five years of standardized spotlight surveys of four segments of a large river in northern Australia revealed that the density of freshwater crocodiles decreased following toad invasion and continued to decline thereafter. Overall, intermediate‐sized freshwater crocodiles (0.6–1.2 m) were most severely impacted. Densities of saltwater crocodiles Crocodylus porosus increased over time and were generally less affected by toad arrival, although toad impacts were inconsistent across survey sections and size classes. Across the entire river, total freshwater crocodile densities declined by 69.5% between 1997 and 2013. Assessments of the status of this species within other large river systems in northern Australia, where baseline data are available from before the toads arrived, should be prioritized. Our findings highlight the importance of long‐term monitoring programmes for quantifying the impacts of novel and unforeseen threats.

Managing Conflicts with Wildlife: Living with Frogs

Frogs control garden pests such as insects and slugs, and serve as a food source for many larger wildlife species. Research on the substances frogs secrete through their skins has even led to the creation of new painkillers and antibiotics. Most frogs in Florida are reclusive and harmless to people, but two species of frogs that have invaded Florida can be harmful to humans and their pets. This 4-page fact sheet presents some facts about native frogs, describes the problems invasive frogs cause, and provides suggestions on how to cope with problem frogs. Written by Steve Johnson, Holly K. Ober, and William M. Giuliano, and published by the UF Department of Wildlife Ecology and Conservation, November 2014. (Photos: Steve Johnson, UF/IFAS) WEC349/UW394: Managing Conflicts with Wildlife: Living with Frogs (ufl.edu)