Femoral bone structure and mechanics at the edge and core of an expanding population of the invasive frog Xenopus laevis.

Understanding how living tissues respond to changes in their mechanical environment is a key question in evolutionary biology. Invasive species provide an ideal model for this as they are often transplanted between environments that differ drastically in their ecological and environmental context. Spatial sorting, the name given to the phenomenon driving differences between individuals at the core and edge of an expanding range, has been demonstrated to impact the morphology and physiology of Xenopus laevis from the invasive French population. Here, we combined a structural analysis using micro-CT scanning and a functional analysis by testing the mechanical properties of the femur to test whether the increased dispersal at the range edge drives differences in bone morphology and function. Our results show significant differences in the inner structure of the femur as well as bone material properties, with frogs from the centre of the range having more robust and resistant bones. This is suggestive of an energy allocation trade-off between locomotion and investment in bone formation, or alternatively, may point to selection for fast locomotion at the range edge. Overall, our results provide insights on the growth of the long bones and the formation of trabecular bone in frogs.

Do muscle contractile properties drive differences in locomotor performance in invasive populations of Xenopus laevis in France?

Jumping and swimming are key locomotor traits in frogs intimately linked to survival and dispersal. French populations of the frog Xenopus laevis from the invasion front are known to possess greater terrestrial locomotor endurance. Here, we tested whether individuals from the invasion front show differences in their muscle physiology that may underlie the observed whole-organism performance differences. We measured muscle contractile properties of the isolated gastrocnemius muscle in vitro, including isometric stress, activation and relaxation time, and work loop power output, both before and during a period of fatiguing contractions. We found that frogs from the centre of the range can produce tetanus force in their gastrocnemius muscle faster than animals from the periphery of the range, which could contribute to higher performance in one-off jumps. Yet, populations did not differ in muscle endurance. These results, coupled with previous work on this invasive population of Xenopus laevis, suggest that the greater stamina observed in individuals from the periphery may be more due to anatomical differences such as longer hind limbs and larger hearts along with potentially other as of yet untested physiological differences rather than differences in the mechanical properties of skeletal muscle.

Rapid Shifts in the Temperature Dependence of Locomotor Performance in an Invasive Frog, Xenopus laevis, Implications for Conservation.

Temperature is a critical abiotic factor impacting all aspects of the biology of organisms, especially in ectotherms. As such, it is an important determinant of the potential invasive ability of organisms and may limit population expansion unless organisms can physiologically respond to changes in temperature either through plasticity or by adapting to their novel environment. Here, we studied the African clawed frog, Xenopus laevis, which has become invasive on a global scale. We compared adults from an invasive population of western France with individuals from two populations in the native range in South Africa. We measured the thermal dependence of locomotor performance in adults given its relevance to dispersal, predator escape, and prey capture. Our results show significant differences in the limits of the 80% performance breadth interval for endurance with the French population showing a left shift in its limits congruent with the colder climate experienced in France. The French invasive population was introduced only about 40 years ago suggesting a rapid shift in the thermal physiology. Given that all individuals were acclimated under laboratory conditions at 23°C for 2 months this suggests that the invasive frogs have adapted to their new environment. These data may allow the refinement of physiologically informed species distribution models permitting better estimates of future ranges at risk of invasion.

Allocation trade-offs impact organ size and muscle architecture in an invasive population of Xenopus laevis in Western France.

Invasive species are a global scourge and often negatively impact native species. Understanding the expansion and dispersal limits of these species is essential. As previous studies have demonstrated increased locomotor performance for populations at the edge of the range of expanding populations, studies of locomotion including the anatomical and physiological traits underlying dispersal capacity are of interest. We focus here on an invasive population of Xenopus laevis introduced in France nearly forty years ago. Previous studies have demonstrated differences in mobility between populations from the centre and the edge of the invasive range, with individuals from the range edge possessing a higher endurance capacity. We test here whether range-edge frogs show anatomical differences in organs or muscles underlying these observed differences of performance. We dissected 10 males and 10 females from central and range-edge sites (40 animals in total) and measured the mass of their organs and the mass, the length, and the physiological cross-sectional area (PCSA) of 28 hind limb muscles. Our results show anatomical differences with individuals from the range edge possessing heavier, longer and more forceful muscles. Moreover, females from the range edge had a heavier heart but lighter stomach than those of the centre of the range. Future studies comparing the morphology between native and invasive populations in other regions or for other species will be especially insightful to better understand the possible adaptive changes in invasive populations and the limits on dispersal capacity.

Acclimation temperature effects on locomotor traits in adult aquatic anurans (X. tropicalis and X. laevis) from different latitudes: possible implications for climate change.

Climate change is in part responsible for the 70% decline in amphibian species numbers worldwide. Although temperature is expected to impact whole-organism performance in ectotherms, reversible thermal acclimation has been suggested as a mechanism that may buffer responses to abrupt temperature changes. Here, we test for an effect of acclimation on locomotor performance traits (jump force and stamina) in adults of two predominantly aquatic and closely related frog species from different climatic regions, Xenopus tropicalis (tropical) and Xenopus laevis (temperate). We find significant effects of acclimation temperature on exertion capacity and for jump force in X. tropicalis but no effect of acclimation temperature on burst performance in X. laevis. Our results suggest that the two locomotor performance traits measured are differentially impacted by acclimation temperature in X. tropicalis. Our results further support the hypothesis that lower-latitude ectotherms might have greater acclimation capacity than high-latitude ones. Finally, our results highlight the importance of investigating multiple performance traits when evaluating how animals may cope with changes in temperature. Further work is required to evaluate the potential for acclimation in mitigating the negative impacts of climate change on amphibian populations.

Global realized niche divergence in the African clawed frog Xenopus laevis.

Although of crucial importance for invasion biology and impact assessments of climate change, it remains widely unknown how species cope with and adapt to environmental conditions beyond their currently realized climatic niches (i.e., those climatic conditions existing populations are exposed to). The African clawed frog Xenopus laevis, native to southern Africa, has established numerous invasive populations on multiple continents making it a pertinent model organism to study environmental niche dynamics. In this study, we assess whether the realized niches of the invasive populations in Europe, South, and North America represent subsets of the species' realized niche in its native distributional range or if niche shifts are traceable. If shifts are traceable, we ask whether the realized niches of invasive populations still contain signatures of the niche of source populations what could indicate local adaptations. Univariate comparisons among bioclimatic conditions at native and invaded ranges revealed the invasive populations to be nested within the variable range of the native population. However, at the same time, invasive populations are well differentiated in multidimensional niche space as quantified via n-dimensional hypervolumes. The most deviant invasive population are those from Europe. Our results suggest varying degrees of realized niche shifts, which are mainly driven by temperature related variables. The crosswise projection of the hypervolumes that were trained in invaded ranges revealed the south-western Cape region as likely area of origin for all invasive populations, which is largely congruent with DNA sequence data and suggests a gradual exploration of novel climate space in invasive populations.

Are invasive populations characterized by a broader diet than native populations?

BACKGROUND: Invasive species are among the most significant threats to biodiversity. The diet of invasive animal populations is a crucial factor that must be considered in the context of biological invasions. A broad dietary spectrum is a frequently cited characteristic of invasive species, allowing them to thrive in a wide range of environments. Therefore, empirical studies comparing diet in invasive and native populations are necessary to understand dietary requirements, dietary flexibility, and the associated impacts of invasive species. METHODS: In this study, we compared the diet of populations of the African clawed frog, Xenopus laevis in its native range, with several areas where it has become invasive. Each prey category detected in stomach contents was assigned to an ecological category, allowing a comparison of the diversity of ecological traits among the prey items in the diet of native and introduced populations. The comparison of diets was also performed using evenness as a niche breadth index on all sampled populations, and electivity as a prey selection index for three out of the six sampled populations. RESULTS: Our results showed that diet breadth could be either narrow or broad in invasive populations. According to diet and prey availability, zooplankton was strongly preferred in most cases. In lotic environments, zooplankton was replaced by benthic preys, such as ephemeropteran larvae. DISCUSSION: The relative proportions of prey with different ecological traits, and dietary variability within and between areas of occurrence, suggest that X. laevis is a generalist predator in both native and invasive populations. Shifts in the realized trophic niche are observed, and appear related to resource availability. Xenopus laevis may strongly impact aquatic ecosystems because of its near complete aquatic lifestyle and its significant consumption of key taxa for the trophic relationships in ponds.

Differences in mobility at the range edge of an expanding invasive population of Xenopus laevis in the west of France.

Theoretical models predict that spatial sorting at the range edge of expanding populations should favor individuals with increased mobility relative to individuals at the center of the range. Despite the fact that empirical evidence for the evolution of locomotor performance at the range edge is rare, data on cane toads support this model. However, whether this can be generalized to other species remains largely unknown. Here, we provide data on locomotor stamina and limb morphology in individuals from two sites: one from the center and one from the periphery of an expanding population of the clawed frog Xenopus laevis in France where it was introduced about 30 years ago. Additionally, we provide data on the morphology of frogs from two additional sites to test whether the observed differences can be generalized across the range of this species in France. Given the known sexual size dimorphism in this species, we also test for differences between the sexes in locomotor performance and morphology. Our results show significant sexual dimorphism in stamina and morphology, with males having longer legs and greater stamina than females. Moreover, in accordance with the predictions from theoretical models, individuals from the range edge had a greater stamina. This difference in locomotor performance is likely to be driven by the significantly longer limb segments observed in animals in both sites sampled in different areas along the range edge. Our data have implications for conservation because spatial sorting on the range edge may lead to an accelerated increase in the spread of this invasive species in France.

Impacts of Climate Change on the Global Invasion Potential of the African Clawed Frog Xenopus laevis.

By altering or eliminating delicate ecological relationships, non-indigenous species are considered a major threat to biodiversity, as well as a driver of environmental change. Global climate change affects ecosystems and ecological communities, leading to changes in the phenology, geographic ranges, or population abundance of several species. Thus, predicting the impacts of global climate change on the current and future distribution of invasive species is an important subject in macroecological studies. The African clawed frog (Xenopus laevis), native to South Africa, possesses a strong invasion potential and populations have become established in numerous countries across four continents. The global invasion potential of X. laevis was assessed using correlative species distribution models (SDMs). SDMs were computed based on a comprehensive set of occurrence records covering South Africa, North America, South America and Europe and a set of nine environmental predictors. Models were built using both a maximum entropy model and an ensemble approach integrating eight algorithms. The future occurrence probabilities for X. laevis were subsequently computed using bioclimatic variables for 2070 following four different IPCC scenarios. Despite minor differences between the statistical approaches, both SDMs predict the future potential distribution of X. laevis, on a global scale, to decrease across all climate change scenarios. On a continental scale, both SDMs predict decreasing potential distributions in the species' native range in South Africa, as well as in the invaded areas in North and South America, and in Australia where the species has not been introduced. In contrast, both SDMs predict the potential range size to expand in Europe. Our results suggest that all probability classes will be equally affected by climate change. New regional conditions may promote new invasions or the spread of established invasive populations, especially in France and Great Britain.

Unequal contribution of native South African phylogeographic lineages to the invasion of the African clawed frog, Xenopus laevis, in Europe.

Due to both deliberate and accidental introductions, invasive African Clawed Frog (Xenopus laevis) populations have become established worldwide. In this study, we investigate the geographic origins of invasive X. laevis populations in France and Portugal using the phylogeographic structure of X. laevis in its native South African range. In total, 80 individuals from the whole area known to be invaded in France and Portugal were analysed for two mitochondrial and three nuclear genes, allowing a comparison with 185 specimens from the native range. Our results show that native phylogeographic lineages have contributed differently to invasive European X. laevis populations. In Portugal, genetic and historical data suggest a single colonization event involving a small number of individuals from the south-western Cape region in South Africa. In contrast, French invasive X. laevis encompass two distinct native phylogeographic lineages, i.e., one from the south-western Cape region and one from the northern regions of South Africa. The French X. laevis population is the first example of a X. laevis invasion involving multiple lineages. Moreover, the lack of population structure based on nuclear DNA suggests a potential role for admixture within the invasive French population.