No evidence for cold-adapted life-history traits in cool-climate populations of invasive cane toads (Rhinella marina).

As an invasive organism spreads into a novel environment, it may encounter strong selective pressures to adapt to abiotic and biotic challenges. We examined the effect of water temperature during larval life on rates of survival and growth of the early life-history stages of cane toads (Rhinella marina) from two geographic regions (tropical vs. temperate) in the species' invaded range in eastern Australia. If local adaptation at the southern (cool-climate) invasion front has extended the cold-tolerance of early life-stages, we would expect to see higher viability of southern-population toads under cooler conditions. Our comparisons revealed no such divergence: the effects of water temperature on rates of larval survival and growth, time to metamorphosis, size at metamorphosis and locomotor performance of metamorphs were similar in both sets of populations. In two cases where tropical and temperate-zone populations diverged in responses to temperature, the tropical animals performed better at low to medium temperatures than did conspecifics from cooler regions. Adaptation to low temperatures in the south might be constrained by behavioural shifts (e.g., in reproductive seasonality, spawning-site selection) that allow toads to breed in warmer water even in cool climates, by gene flow from warmer-climate populations, or by phylogenetic conservatism in these traits.

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.

Moving south: effects of water temperatures on the larval development of invasive cane toads (Rhinella marina) in cool-temperate Australia.

The distributional limits of many ectothermic species are set by thermal tolerances of early-developmental stages in the life history; embryos and larvae often are less able to buffer environmental variation than are conspecific adults. In pond-breeding amphibians, for example, cold water may constrain viability of eggs and larvae, even if adults can find suitable thermal conditions in terrestrial niches. Invasive species provide robust model systems for exploring these questions, because we can quantify thermal challenges at the expanding range edge (from field surveys) and larval responses to thermal conditions (in the laboratory). Our studies on invasive cane toads (Rhinella marina) at the southern (cool-climate) edge of their expanding range in Australia show that available ponds often average around 20°C during the breeding period, 10°C lower than in many areas of the toads' native range, or in the Australian tropics. Our laboratory experiments showed that cane toad eggs and larvae cannot develop successfully at 16°C, but hatching success and larval survival rates were higher at 20°C than in warmer conditions. Lower temperatures slowed growth rates, increasing the duration of tadpole life, but also increased metamorph body mass. Water temperature also influenced metamorph body shape (high temperatures reduced relative limb length, head width, and body mass) and locomotor performance (increased speed from intermediate temperatures, longer hops from high temperatures). In combination with previous studies, our data suggest that lower water temperatures may enhance rather than reduce recruitment of cane toads, at least in areas where pond temperatures reach or exceed 20°C. That condition is fulfilled over a wide area of southern Australia, suggesting that the continuing expansion of this invasive species is unlikely to be curtailed by the impacts of relatively low water temperatures on the viability of early life-history stages.

The Acid Test: pH Tolerance of the Eggs and Larvae of the Invasive Cane Toad (Rhinella marina) in Southeastern Australia.

Invasive cane toads are colonizing southeastern Australia via a narrow coastal strip sandwiched between unsuitable areas (Pacific Ocean to the east, mountains to the west). Many of the available spawning sites exhibit abiotic conditions (e.g., temperature, salinity, and pH) more extreme than those encountered elsewhere in the toad's native or already invaded range. Will that challenge impede toad expansion? To answer that question, we measured pH in 35 ponds in northeastern New South Wales and 8 ponds in the Sydney region, in both areas where toads occur (and breed) and adjacent areas where toads are likely to invade, and conducted laboratory experiments to quantify effects of pH on the survival and development of toad eggs and larvae. Our field surveys revealed wide variation in pH (3.9-9.8) among natural water bodies. In the laboratory, the hatching success of eggs was increased at low pH (down to pH 4), whereas the survival, growth, and developmental rates of tadpoles were enhanced by higher pH levels. We found that pH influenced metamorph size and shape (relative head width, relative leg length) but not locomotor performance. The broad tolerance range of these early life-history stages suggests that pH conditions in ponds will not significantly slow the toad's expansion southward. Indeed, toads may benefit from transiently low pH conditions, and habitat where pH in wetlands is consistently low (such as coastal heath) may enhance rather than reduce toad reproductive success. A broad physiological tolerance during embryonic and larval life has contributed significantly to the cane toad's success as a widespread colonizer.