Sex-based differences in the use of post-fire habitats by invasive cane toads (Rhinella marina).

Wildfires can modify habitat attributes, and those changes may differentially affect males versus females within a species if there is pre-existing niche divergence between the sexes. We used radio-tracking and dissections to study invasive cane toads (Rhinella marina), and performed transect counts on native frogs and cane toads 12 months after extensive fires in forests of eastern Australia. Both toads and native frogs were encountered more frequently in burned sites than in unburned sites. Most microhabitat features were similar between burned versus unburned areas, but fire had differential impacts on the ecology of male versus female toads. In burned areas females were less numerous but were larger, in better body condition, and had consumed more prey (especially, coleopterans and myriapods). The impact of fire on attributes of retreat-sites (e.g., temperature, density of vegetation cover) also differed between the sexes. More generally, intraspecific divergence in ecological traits within a species (as a function of body size as well as sex) may translate into substantial divergences in the impacts of habitat change.

Wildfires modify the parasite loads of invasive cane toads.

The frequency and severity of wildfires are increasing due to anthropogenic modifications to habitats and to climate. Post-fire landscapes may advantage invasive species via multiple mechanisms, including changes to host-parasite interactions. We surveyed the incidence of endoparasitic lungworms (Rhabdias pseudosphaerocephala) in invasive cane toads (Rhinella marina) in near-coastal sites of eastern Australia, a year after extensive fires in this region. Both the prevalence of infection and number of worms in infected toads increased with toad body size in unburned areas. By contrast, parasite load decreased with toad body size in burned areas. By killing moisture-dependent free-living lungworm larvae, the intense fires may have liberated adult cane toads from a parasite that can substantially reduce the viability of its host. Smaller toads, which are restricted to moist environments, did not receive this benefit from fires.

Physiological plasticity in a successful invader: rapid acclimation to cold occurs only in cool-climate populations of cane toads (Rhinella marina).

Physiological plasticity may facilitate invasion of novel habitats; but is such plasticity present in all populations of the invader or is it elicited only by specific climatic challenges? In cold-climate areas of Australia, invasive cane toads (Rhinella marina) can rapidly acclimate to cool conditions. To investigate whether this physiological plasticity is found in all invasive cane toads or is only seen in cool climates, we measured the acclimation ability of toads from across Australia and the island of Hawai'i. We collected toads from the field and placed them at either 12 or 24°C for 12 h before measuring their righting response as a proxy for critical thermal minimum (CTmin). Toads from the coolest Australian region (New South Wales) demonstrated plasticity (as previously reported), with exposure to 12°C (vs. 24°C) decreasing CTmin by 2°C. In toads from other Australian populations, CTmins were unaffected by our thermal treatments. Hawai'ian toads from a cool, wet site also rapidly acclimated to cool conditions, whereas those from warmer and drier Hawai'ian sites did not. Thermal plasticity has diverged among populations of invasive cane toads, with rapid acclimation manifested only in two cool-climate populations from widely separated sites. Predictions about the potential range of invasive species thus must consider the possibility of geographic (intraspecific) heterogeneity in thermal plasticity; data from other parts of the species' range may fail to predict levels of plasticity elicited by thermal challenges.

New Weapons in the Toad Toolkit: A Review of Methods to Control and Mitigate the Biodiversity Impacts of Invasive Cane Toads (Rhinella Marina).

Our best hope of developing innovative methods to combat invasive species is likely to come from the study of high-profile invaders that have attracted intensive research not only into control, but also basic biology. Here we illustrate that point by reviewing current thinking about novel ways to control one of the world's most well-studied invasions: that of the cane toad in Australia. Recently developed methods for population suppression include more effective traps based on the toad's acoustic and pheromonal biology. New tools for containing spread include surveillance technologies (e.g., eDNA sampling and automated call detectors), as well as landscape-level barriers that exploit the toad's vulnerability to desiccation­a strategy that could be significantly enhanced through the introduction of sedentary, range-core genotypes ahead of the invasion front. New methods to reduce the ecological impacts of toads include conditioned taste aversion in free-ranging predators, gene banking, and targeted gene flow. Lastly, recent advances in gene editing and gene drive technology hold the promise of modifying toad phenotypes in ways that may facilitate control or buffer impact. Synergies between these approaches hold great promise for novel and more effective means to combat the toad invasion and its consequent impacts on biodiversity.

Is the enhanced dispersal rate seen at invasion fronts a behaviourally plastic response to encountering novel ecological conditions?

As a population expands into novel areas (as occurs in biological invasions), the range edge becomes dominated by rapidly dispersing individuals-thereby accelerating the rate of population spread. That acceleration has been attributed to evolutionary processes (natural selection and spatial sorting), to which we add a third complementary process: behavioural plasticity. Encountering environmental novelty may directly elicit an increased rate of dispersal. When we reciprocally translocated cane toads (Rhinella marina) among study sites in southern Australia, the transported animals massively increased dispersal rates relative to residents (to an extent similar to the evolved increase between range-core versus invasion-front toad populations in Australia). The responses of these translocated toads show that even range-core toads are capable of the long-distance dispersal rates of invasion-front conspecifics and suggest that rapid dispersal (rather than evolving de novo) has simply been expanded from facultative to constitutive expression.

Toads on Lava: Spatial Ecology and Habitat Use of Invasive Cane Toads (Rhinella marina) in Hawai'i.

Most ecological research on cane toads (Rhinella marina) has focused on invasive populations in Australia, ignoring other areas where toads have been introduced. We radio-tracked and spool-tracked 40 toads, from four populations on the island of Hawai'i. Toads moved extensively at night (mean 116 m, from spool-tracking) but returned to the same or a nearby retreat-site each day (from radio-tracking, mean distance between successive retreat sites 11 m; 0 m for 70% of records). Males followed straighter paths during nocturnal movements than did females. Because moist sites are scarce on the highly porous lava substrate, Hawai'ian toads depend on anthropogenic disturbance for shelter (e.g. beneath buildings), foraging (e.g. suburban lawns, golf courses) and breeding (artificial ponds). Foraging sites are further concentrated by a scarcity of flying insects (negating artificial lights as prey-attractors). Habitat use of toads shifted with time (at night, toads selected areas with less bare ground, canopy, understory and leaf-litter), and differed between sexes (females foraged in areas of bare ground with dense understory vegetation). Cane toads in Hawai'i thrive in scattered moist patches within a severely arid matrix, despite a scarcity of flying insects, testifying to the species' ability to exploit anthropogenic disturbance.

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.

Do invasive cane toads affect the parasite burdens of native Australian frogs?

One of the most devastating impacts of an invasive species is the introduction of novel parasites or diseases to native fauna. Invasive cane toads (Rhinella marina) in Australia contain several types of parasites, raising concern that the toads may increase rates of parasitism in local anuran species. We sampled cane toads and sympatric native frogs (Limnodynastes peronii, Litoria latopalmata, and Litoria nasuta) at the southern invasion front of cane toads in north-eastern New South Wales (NSW). We dissected and swabbed these anurans to score the presence and abundance of nematodes (Rhabdias lungworms, and gastric encysting nematodes), myxozoans, and chytrid fungus. To determine if cane toad invasion influences rates of parasitism in native frogs, we compared the prevalence and intensity of parasites in frogs from areas with toads, to frogs from areas without toads. Contrary to the situation on the (rapidly-expanding) tropical invasion front, cane toads on the slowly-expanding southern front were heavily infected with rhabditoid lungworms. Toads also contained gastric-encysting nematodes, and one toad was infected by chytrid fungus, but we did not find myxozoans in any toads. All parasite groups were recorded in native frogs, but were less common in areas invaded by toads than in nearby yet to be invaded areas. Contrary to our predictions, toad invasion was associated with a reduced parasite burden in native frogs. Thus, cane toads do not appear to transfer novel parasites to native frog populations, or act as a reservoir for native parasites to 'spill-back' into native frogs. Instead, cane toads may reduce frog-parasite numbers by taking up native parasites that are then killed by the toad's immune defences.