ALBERT

Description: Like most ectothermic vertebrates, keelback snakes ( Tropidonophis mairii ) do not exhibit parental care. Thus, offspring must possess an immune system capable of dealing with challenges such as pathogens, without assistance from an attendant parent. We know very little about immune system characteristics of neonatal reptiles, including the magnitude of heritability and other maternal influences. To identify sources of variation in circulating white blood cell (WBC) concentrations and differentials, we examined blood smears from 246 hatchling snakes and their field-caught mothers. WBC concentrations were lower in hatchlings than in adults, and hatchlings had more basophils and fewer azurophils than adults. A hatchling keelback's WBC differential was also influenced by its sex and body size. Although hatchling WBC measures exhibited negligible heritability, they were strongly influenced by maternal body size and parasite infection (but not by maternal body condition, relative clutch mass or time in captivity). Larger mothers produced offspring with more azurophils and fewer lymphocytes. The mechanisms and consequences of WBC variation are currently unknown, but if these maternal effects enhance offspring fitness, the impact of maternal body size on reproductive success may be greater than expected simply from allometric increases in the numbers and sizes of progeny.

Description: Invasive species provide a robust opportunity to evaluate how animals deal with novel environmental challenges. Shifts in locomotor performance—and thus the ability to disperse—(and especially, the degree to which it is constrained by thermal and hydric extremes) are of special importance, because they might affect the rate that an invader can spread. We studied cane toads ( Rhinella marina ) across a broad geographical range: two populations within the species' native range in Brazil, two invasive populations on the island of Hawai'i and eight invasive populations encompassing the eastern, western and southern limits of the toad invasion in Australia. A toad's locomotor performance on a circular raceway was strongly affected by both its temperature and its hydration state, but the nature and magnitude of those constraints differed across populations. In their native range, cane toads exhibited relatively low performance (even under optimal test conditions) and a rapid decrease in performance at lower temperatures and hydration levels. At the other extreme, performance was high in toads from southern Australia, and virtually unaffected by desiccation. Hawai'ian toads broadly resembled their Brazilian conspecifics, plausibly reflecting similar climatic conditions. The invasion of Australia has been accompanied by a dramatic enhancement in the toads' locomotor abilities, and (in some populations) by an ability to maintain locomotor performance even when the animal is cold and/or dehydrated. The geographical divergences in performance among cane toad populations graphically attest to the adaptability of invasive species in the face of novel abiotic challenges.

Description: Invasive species must deal with novel challenges, both from the alien environment and from pressures arising from range expansion per se (e.g. spatial sorting). Those conditions can create geographical variation in behaviour across the invaded range, as has been documented across regions of Australia invaded by cane toads; range-edge toads are more exploratory and willing to take risks than are conspecifics from the range-core. That behavioural divergence might be a response to range expansion and invasion per se , or to the different environments encountered. Climate differs across the cane toads' invasion range from the wet tropics of Queensland to the seasonally dry climates of northwestern Western Australia. The different thermal and hydric regimes may affect behavioural traits via phenotypic plasticity or through natural selection. We cannot tease apart the effects of range expansion versus climate in an expanding population but can do so in a site where the colonizing species was simultaneously released in all suitable areas, thus removing any subsequent phase of range expansion. Cane toads were introduced to Hawai'i in 1932; and thence to Australia in 1935. Toads were released in all major sugarcane-growing areas in Hawai'i within a 12-month period. Hence, Hawai'ian cane toads provide an opportunity to examine geographical divergence in behavioural traits in a climatically diverse region (each island has both wet and dry sides) in the absence of range expansion subsequent to release. We conducted laboratory-based behavioural trials testing exploration, risk-taking and response to novelty using field-caught toads from the wet and dry sides of two Hawai'ian islands (Oahu and Hawai'i). Toads from the dry side of Oahu had a higher propensity to take risks than did toads from the dry side of Hawai'i. Toads from Oahu were also more exploratory than were conspecifics from the island of Hawai'i. However, toads from wet versus dry climates were similar in all behaviours that we scored, suggesting that founder effects, genetic drift, or developmentally plastic responses to ecological factors other than climate may have driven behavioural divergence between islands.

Description: Individuals at the leading edge of expanding biological invasions often show distinctive phenotypic traits, in ways that enhance their ability to disperse rapidly and to function effectively in novel environments. Cane toads ( Rhinella marina ) at the invasion front in Australia exhibit shifts in morphology, physiology and behaviour (directionality of dispersal, boldness, risk-taking). We took a common-garden approach, raising toads from range-core and range-edge populations in captivity, to see if the behavioural divergences observed in wild-caught toads are also evident in common-garden offspring. Captive-raised toads from the invasion vanguard population were more exploratory and bolder (more prone to ‘risky’ behaviours) than toads from the range core, which suggests that these are evolved, genetic traits. Our study highlights the importance of behaviour as being potentially adaptive in invasive populations and adds these behavioural traits to the increasing list of phenotypic traits that have evolved rapidly during the toads' 80-year spread through tropical Australia.

Description: Invasive species often exhibit rapid evolutionary changes, and can provide powerful insights into the selective forces shaping phenotypic traits that influence dispersal rates and/or sexual interactions. Invasions also may modify sexual dimorphism. We measured relative lengths of forelimbs and hindlimbs of more than 3000 field-caught adult cane toads ( Rhinella marina ) from 67 sites in Hawai'i and Australia (1–80 years post-colonization), along with 489 captive-bred individuals from multiple Australian sites raised in a ‘common garden’ (to examine heritability and reduce environmental influences on morphology). As cane toads spread from east to west across Australia, the ancestral condition (long limbs, especially in males) was modified. Limb length relative to body size was first reduced (perhaps owing to natural selection on locomotor ability), but then increased again (perhaps owing to spatial sorting) in the invasion vanguard. In contrast, the sex disparity in relative limb length has progressively decreased during the toads' Australian invasion. Offspring reared in a common environment exhibited similar geographical divergences in morphology as did wild-caught animals, suggesting a genetic basis to the changes. Limb dimensions showed significant heritability (2–17%), consistent with the possibility of an evolved response. Cane toad populations thus have undergone a major shift in sexual dimorphism in relative limb lengths during their brief (81 years) spread through tropical Australia.