A trophic cascade initiated by an invasive vertebrate alters the structure of native reptile communities

Invasive vertebrates are frequently reported to have catastrophic effects on the populations of species which they directly impact. It follows then, that if invaders exert strong suppressive effects on some species then other species will indirectly benefit due to ecological release from interactions with directly impacted species. However, evidence that invasive vertebrates trigger such trophic cascades and alter community structure in terrestrial ecosystems remains rare. Here, we ask how the cane toad, a vertebrate invader that is toxic to many of Australia's vertebrate predators, influences lizard assemblages in a semi‐arid rangeland. In our study area, the density of cane toads is influenced by the availability of water accessible to toads. We compared an index of the abundance of sand goannas, a large predatory lizard that is susceptible to poisoning by cane toads and the abundances of four lizard families preyed upon by goannas (skinks, pygopods, agamid lizards and geckos) in areas where cane toads were common or rare. Consistent with the idea that suppression of sand goannas by cane toads initiates a trophic cascade, goanna activity was lower and small lizards were more abundant where toads were common. The hypothesis that suppression of sand goannas by cane toads triggers a trophic cascade was further supported by our findings that small terrestrial lizards that are frequently preyed upon by goannas were more affected by toad abundance than arboreal geckos, which are rarely consumed by goannas. Furthermore, the abundance of at least one genus of terrestrial skinks benefitted from allogenic ecosystem engineering by goannas where toads were rare. Overall, our study provides evidence that the invasion of ecosystems by non‐native species can have important effects on the structure and integrity of native communities extending beyond their often most obvious and frequently documented direct ecological effects.     

Evolution of dispersal and life history interact to drive accelerating spread of an invasive species

Populations on the edge of an expanding range are subject to unique evolutionary pressures acting on their life‐history and dispersal traits. Empirical evidence and theory suggest that traits there can evolve rapidly enough to interact with ecological dynamics, potentially giving rise to accelerating spread. Nevertheless, which of several evolutionary mechanisms drive this interaction between evolution and spread remains an open question. We propose an integrated theoretical framework for partitioning the contributions of different evolutionary mechanisms to accelerating spread, and we apply this model to invasive cane toads in northern Australia. In doing so, we identify a previously unrecognised evolutionary process that involves an interaction between life‐history and dispersal evolution during range shift. In roughly equal parts, life‐history evolution, dispersal evolution and their interaction led to a doubling of distance spread by cane toads in our model, highlighting the potential importance of multiple evolutionary processes in the dynamics of range expansion.     

The extra-limital spread of an invasive species via 'stowaway' dispersal: toad to nowhere?

The mechanisms by which invasive species spread through new areas can influence the spatial scale of their impact. Although previous research has focused on 'natural' dispersal rates following initial introductions, human-aided inadvertent dispersal by 'stowaways' on commercial and domestic transport is thought to be a major contributor to long-distance dispersal. Few data exist to support this assumption. Cane toads Bufo marinus were introduced to north-eastern Australia in 1935, and have since dispersed rapidly through the tropics. Based on information accumulated by community groups in Sydney, 400 km south of the cane toads' current Australian distribution, we document high rates of translocation (at least 50 toads arriving in Sydney per year). Most toads were translocated on commercial truck transport carrying landscaping and building materials from the current range of the cane toads in New South Wales and Queensland, and resulted in highly clumped locations of toad arrival reflecting primary truck transport destinations. Most introductions involved single toads (68 of 102 translocation events), but some introductions involved two to 19 animals. Adults of both sexes were represented equally but juveniles were rarely detected. High rates of translocation of adult toads of both sexes suggest that the eventual distribution of cane toads in Australia may be limited by the animals' bioclimatic tolerances rather than by an inability to reach suitable habitats, even in areas far distant from the toads' current range.     

High evolutionary constraints limited adaptive responses to past climate changes in toad skulls

Interactions among traits that build a complex structure may be represented as genetic covariation and correlation. Genetic correlations may act as constraints, deflecting the evolutionary response from the direction of natural selection. We investigated the relative importance of drift, selection, and constraints in driving skull divergence in a group of related toad species. The distributional range of these species encompasses very distinct habitats with important climatic differences and the species are primarily distinguished by differences in their skulls. Some parts of the toad skull, such as the snout, may have functional relevance in reproductive ecology, detecting water cues. Thus, we hypothesized that the species skull divergence was driven by natural selection associated with climatic variation. However, given that all species present high correlations among skull traits, our second prediction was of high constraints deflecting the response to selection. We first extracted the main morphological direction that is expected to be subjected to selection by using within- and between-species covariance matrices. We then used evolutionary regressions to investigate whether divergence along this direction is explained by climatic variation between species. We also used quantitative genetics models to test for a role of random drift versus natural selection in skull divergence and to reconstruct selection gradients along species phylogeny. Climatic variables explained high proportions of between-species variation in the most selected axis. However, most evolutionary responses were not in the direction of selection, but aligned with the direction of allometric size, the dimension of highest phenotypic variance in the ancestral population. We conclude that toad species have responded to selection related to climate in their skulls, yet high evolutionary constraints dominated species divergence and may limit species responses to future climate change.     

Seasonal changes in the thermoregulatory strategies of Rhinella arenarum in the Monte desert, Argentina

We determined and compared the efficiency of thermoregulation of Rhinella arenarum in the Monte desert (Argentina) in two seasons, dry and wet. In the field, we measured body temperatures, micro-habitat temperatures and operative temperatures; while in the laboratory; we measured the selected body temperatures. Our results show a change in the thermoregulatory strategy of R. arenarum that is related to environmental constraints on their thermal niche. R. arenarum has the ability to be plastic and combine two strategies: (i) a moderate thermoregulator in the wet season, where thermal resources are available such that body temperature is maintained within the set points and physiologicial and behavioral processes are optimized; and (ii) a thermoconformer in the dry season where the thermal environment is more homogeneous and there is greater time invested in searching for food.     

Barriers,rather than refugia,underlie the origin of diversity in toads endemic to the Brazilian Atlantic Forest

In this study, we investigated the relative contribution of geographic barriers and Pleistocene refuges in the diversification of the Rhinella crucifer species complex, a group of endemic toads with a widespread distribution in the Brazilian Atlantic Forest (AF). We used intensive sampling and multilocus DNA sequence data to compare nucleotide diversity between refuge and nonrefuge areas, investigate regional demographic patterns, estimate demographic parameters related to genetic breaks and test refuge versus barrier scenarios of diversification using approximate Bayesian computation. We did not find higher levels of genetic diversity in putative refuge areas, either at regional or biome scale. Rather, the demographic history of the species complex supports regional differences with moderate population growth in the north and central regions and stability in southern AF. Genetic breaks were dated to the Plio–Pleistocene; however, our analyses rejected the role of refuges in creating a northern and central divergence, supporting a recent colonization scenario at a smaller scale within the central AF. Overall, our data rule out massive climatically driven fragmentation and large‐scale recolonization events for populations across the biome. We confirmed the importance of geographic barriers in creating main divergences and underscored the importance of searching for cryptic discontinuities in the landscape. Comparison of our results with those of other AF taxa indicates organismal specific responses to moderate shifts in habitat and that multiple refuges may constitute a more realistic model for diversification of Atlantic Forest biota.     

The thermal dependency of locomotor performance evolves rapidly within an invasive species

Biological invasions can stimulate rapid shifts in organismal performance, via both plasticity and adaptation. We can distinguish between these two proximate mechanisms by rearing offspring from populations under identical conditions and measuring their locomotor abilities in standardized trials. We collected adult cane toads (Rhinella marina) from invasive populations that inhabit regions of Australia with different climatic conditions. We bred those toads and raised their offspring under common‐garden conditions before testing their locomotor performance. At high (but not low) temperatures, offspring of individuals from a hotter location (northwestern Australia) outperformed offspring of conspecifics from a cooler location (northeastern Australia). This disparity indicates that, within less than 100 years, thermal performance in cane toads has adapted to the novel abiotic challenges that cane toads have encountered during their invasion of tropical Australia.     

Differences in developmental strategies between long‐settled and invasion‐front populations of the cane toad in Australia

Phenotypic plasticity can enhance a species’ ability to persist in a new and stressful environment, so that reaction norms are expected to evolve as organisms encounter novel environments. Biological invasions provide a robust system to investigate such changes. We measured the rates of early growth and development in tadpoles of invasive cane toads (Rhinella marina) in Australia, from a range of locations and at different larval densities. Populations in long‐colonized areas have had the opportunity to adapt to local conditions, whereas at the expanding range edge, the invader is likely to encounter challenges that are both novel and unpredictable. We thus expected invasion‐vanguard populations to exhibit less phenotypic plasticity than range‐core populations. Compared to clutches from long‐colonized areas, clutches from the invasion front were indeed less plastic (i.e. rates of larval growth and development were less sensitive to density). In contrast, those rates were highly variable in clutches from the invasion front, even among siblings from the same clutch under standard conditions. Clutches with highly variable rates of growth and development under constant conditions had lower phenotypic plasticity, suggesting a trade‐off between these two strategies. Although these results reveal a strong pattern, further investigation is needed to determine whether these different developmental strategies are adaptive (i.e. adaptive phenotypic plasticity vs. bet‐hedging) or instead are driven by geographic variation in genetic quality or parental effects.