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Abstract Estimating the size of animal populations plays an important role in evidence‐based conservation and management. Some methods for estimating population size rely on animals being individually identifiable. Traditionally, this has been done by marking physically captured animals, but increasingly, animals with distinctive natural markings are surveyed noninvasively using cameras. Animal reidentification from photographs is usually done manually, which is expensive, laborious, and requires considerable skill. An alternative is to develop computer vision methods that can support or replace the manual identification task. We developed an automated approach using deep learning to identify whether a pair of photographs is of the same individual or not. The core of the approach is a similarity learning network that uses paired convolutional neural networks with a triplet loss function to summarize image pairs and decide whether they are from the same individual. Prior to the main matching step, two additional convolutional neural networks perform image segmentation, cropping the animal object within the image, and orientation prediction, deciding which side of the animal was photographed. We applied the approach to four species, with images of the same individual often spanning several years: systematic surveys of bottlenose dolphins (Tursiops truncatus, 2008–2019) and harbor seals (Phoca vitulina, 2015–2019), a citizen science dataset of western leopard toads (Sclerophrys pantherina, unknown dates), and a publicly available repository of humpback whale images (Megaptera novaeangliae, unknown dates). For these species, our best‐performing models were able to identify whether a pair of images were from the same individual or different individuals in 95.8%, 94.6%, 88.2%, and 83.8% of the cases, respectively. We found that triplet loss functions outperformed binary cross‐entropy loss functions and that data augmentation and additional manual curation of training data provided small but consistent improvements in performance. These results demonstrate the potential of deep learning to replace or, more likely, support and facilitate manual individual identification efforts.

Abstract Sub-Saharan Africa is under-represented in global biodiversity datasets, particularly regarding the impact of land use on species’ population abundances. Drawing on recent advances in expert elicitation to ensure data consistency, 200 experts were convened using a modified-Delphi process to estimate ‘intactness scores’: the remaining proportion of an ‘intact’ reference population of a species group in a particular land use, on a scale from 0 (no remaining individuals) to 1 (same abundance as the reference) and, in rare cases, to 2 (populations that thrive in human-modified landscapes). The resulting bii4africa dataset contains intactness scores representing terrestrial vertebrates (tetrapods: ±5,400 amphibians, reptiles, birds, mammals) and vascular plants (±45,000 forbs, graminoids, trees, shrubs) in sub-Saharan Africa across the region’s major land uses (urban, cropland, rangeland, plantation, protected, etc.) and intensities (e.g., large-scale vs smallholder cropland). This dataset was co-produced as part of the Biodiversity Intactness Index for Africa Project. Additional uses include assessing ecosystem condition; rectifying geographic/taxonomic biases in global biodiversity indicators and maps; and informing the Red List of Ecosystems.

Amphibians are more threatened than any other vertebrate class, yet evidence for many threats is missing. The Cape lowland fynbos (endemic scrub biome) is threatened by habitat loss, and natural temporary freshwater habitats are removed in favour of permanent impoundments. In this study, we determine amphibian assemblages across different freshwater habitat types with special attention to the presence of invasive fish. We find that anuran communities differ primarily by habitat type, with permanent water habitats having more widespread taxa, while temporary water bodies have more range restricted taxa. Invasive fish are found to have a significant impact on frogs with toads most tolerant of their presence. Temporary freshwater habitats are a conservation priority in the area, and their amphibian assemblages represent endemic taxa that are intolerant of invasive fish. Conservation of a biodiverse amphibian assemblage in lowland fynbos areas will rely on the creation of temporary freshwater habitats, rather than a northern hemisphere pond based solution.

Angola has experienced an incredible increase of the knowledge of its herpetofauna over the past decade. However, accurate biodiversity inventories remain deficient for certain regions of particular conservation interest. We therefore provide an updated checklist of Iona National Park’s herpetofauna, with 75 recorded species, including five amphibians and 70 reptiles, 40 of these recorded for the first time in Iona National Park. These species comprise ~80% of the reptile diversity of Namibe Province. Therefore, this work recognises Iona National Park as the most reptile-diverse protected area in Angola and is potentially one of the richest in southern Africa. Consequently, this work enhances the importance of specific conservation plans in the area and the need for further investigation into the hidden biodiversity of this region.

Management strategies for invasive populations should be designed to maximise efficacy and efficiency, i.e. to accomplish their goals while operating with the least resource consumption. This optimisation is often difficult to achieve in stage-structured populations, because costs, benefits and feasibility of removing individuals may vary with stage. We use a spatially-explicit stage-structured model to assess efficacy of past, present and alternative control strategies for invasive guttural toads, Sclerophrys gutturalis, in Cape Town. The strategies involve removal of variable proportions of individuals at different life-history stages and spatial scales. We also quantify the time necessary to implement each strategy as a proxy of financial resources and we correct strategy outcomes by implementation of time to estimate efficiency. We found that the strategy initially pursued in Cape Town, which did not target any specific stage, was less efficient than the present strategy, which prioritises adult removal. The initial strategy was particularly inefficient because it did not reduce the population size despite allocating consistent resources to remove eggs and tadpoles. We also found that such removal might be detrimental when applied at high levels. This counter-intuitive outcome is due to the ‘hydra effect’: an undesired increase in population size caused by removing individuals before overcompensatory density dependence. Strategies that exclusively remove adults ensure much greater management efficiency than those that also remove eggs and tadpoles. Available management resources should rather be allocated to increase the proportion of adult guttural toads that are removed or the spatial extent at which this removal is pursued.

AbstractAllotetraploid genomes have two distinct genomic components called subgenomes that are derived from separate diploid ancestral species. Many genomic characteristics such as gene function, expression, recombination, and transposable element mobility may differ significantly between subgenomes. To explore the possibility that subgenome population structure and gene flow may differ as well, we examined genetic variation in an allotetraploid frog—the African clawed frog (Xenopus laevis

Cities are focal points of introduction for invasive species. Urban evolution might facilitate the success of invasive species in recipient urban habitats. Here we test this hypothesis by rearing tadpoles of a successful amphibian urban coloniser and invader in a common garden environment. We compared growth rate, morphological traits, swimming performance, and developmental rate of guttural toad tadpoles (Sclerophrys gutturalis) from native rural, native urban, and non-native urban habitats. By measuring these traits across ontogeny, we were also able to compare divergence across different origins as the tadpoles develop. The tadpoles of non-native urban origin showed significantly slower developmental rate (e.g., the proportion of tadpoles reaching Gosner stage 31 or higher was lower at age 40 days) than tadpoles of native urban origin. Yet, tadpoles did not differ in growth rate or any morphological or performance trait examined, and none of these traits showed divergent ontogenetic changes between tadpoles of different origin. These findings suggest that prior adaptation to urban habitats in larval traits likely does not play an important role in facilitating the invasion success of guttural toads into other urban habitats. Instead, we suggest that evolutionary changes in larval traits after colonization (e.g., developmental rate), together with decoupling of other traits and phenotypic plasticity might explain how this species succeeded in colonising extra-limital urban habitats.

Abstract Estimating and planning for the impacts of climate change on the biodiversity of protected areas is a major challenge for conservation managers. When these areas are topographically heterogenous and contain species' entire ranges, this challenge is exacerbated because the coarse spatial scales of Global Circulation Model projections provide limited information for within‐park management. South Africa's Table Mountain National Park, home to three endemic amphibian species in just ~24,500 hectares, provides a case study for identifying conservation needs under climate change. Selecting the park's herpetofauna as pilot taxa, we identified life history and demographic characteristics believed to make species more sensitive and less able to adapt to climate change. We organized these into assessment frameworks and, through a combination of literature review and expert elicitation, reviewed and used them to assess climate change vulnerability of 18 amphibian and 41 reptile species. The assessment highlighted that 73% and 67% of the park's reptile and amphibian species, respectively, had at least one high‐sensitivity and low‐adaptive capacity trait. Using ordinal and additive scoring methods, we identified the species most vulnerable to climate change and highlight the park areas containing their highest concentrations. These areas will be used to inform landscape‐scale management priorities and park use zones. The current IUCN Red List assessments for these species do not incorporate climate change vulnerability. Considering some species appear to be threatened by climate change, their conservation needs might be underestimated. Identifying the most vulnerable species and the mechanisms underpinning their vulnerability can guide the identification and prioritization of conservation needs, while the highlighted knowledge gaps inform priorities for monitoring and research. While comprehensive climate change adaptation planning for Table Mountain National Park requires additional assessment of other taxonomic groups, this trait‐based assessment example highlights a viable tool for assessing climate change vulnerability in protected areas.

The environmental and socio-economic impacts of invasive species have long been recognised to be unequal, with some species being benign while others are disastrous. Until recently there was no recognised standard impact scoring framework with which to compare impacts of species from very different taxa. The advent of the Environmental Impact Classification for Alien Taxa (EICAT) and Socio‐Economic Impact Classification of Alien Taxa (SEICAT) schemes allows for the possibility of assessing impact through a standard approach. However, both these schemes are still in their infancy and the associated costs of the research that informs them is unknown. We aimed to determine the study costs and complexity associated with assessing invasive species’ socio-economic and environmental impacts. We used amphibians as a model group to investigate papers from which EICAT and SEICAT scores could be drawn up to 2019. Our analysis shows that studies that resulted in higher impact scores were more costly. Furthermore, the costs of studies were best predicted by their complexity and the time taken to complete them. If impact scores from EICAT and SEICAT are allowed to inform policy, then we need to carefully consider whether species with low scores represent true impact, or require more research investment and time. Policy makers needing accurate assessments will need to finance larger, more complex, and rigorous studies. Assessing impacts in low and middle income countries may need investment using international research collaborations and capacity building with scientists from high income areas.

A prominent feature of the modern era is the increasing spread of invasive species, particularly within island and urban ecosystems, and these occurrences provide valuable natural experiments by which evolutionary and invasion hypotheses can be tested. In this study, we used the invasion route of guttural toads (Sclerophrys gutturalis) from natural-native and urban-native populations (Durban, South Africa) to their urban-invasive and natural-invasive populations (Mauritius and Réunion) to determine whether phenotypic changes that arose once the toads became urbanized in their native range have increased their invasive potential before they were transported (i.e., prior adaptation) or whether the observed changes are unique to the invasive populations. This urban/natural by native/invasive gradient allowed us to examine differences in guttural toad morphology (i.e., body size, hindlimb, and hindfoot length) and performance capacity (i.e., escape speed, endurance, and climbing ability) along their invasion route. Our findings indicate that invasive island populations have reduced body sizes, shorter limbs in relation to snout-vent length, decreased escape speeds, and decreased endurance capacities that are distinct from the native mainland populations (i.e., invasion-derived change). Thus, these characteristics did not likely arise directly from a pre-transport anthropogenic “filter” (i.e., urban-derived change). Climbing ability, however, did appear to originate within the urban-native range and was maintained within the invasive populations, thereby suggesting it may have been a prior adaptation that provided this species with an advantage during its establishment in urban areas and spread into natural forests. We discuss how this shift in climbing performance may be ecologically related to the success of urban and invasive guttural toad populations, as well as how it may have impacted other island-derived morphological and performance phenotypes.

Abstract Community and invasion ecology have mostly grown independently. There is substantial overlap in the processes captured by different models in the two fields, and various frameworks have been developed to reduce this redundancy and synthesize information content. Despite broad recognition that community and invasion ecology are interconnected, a process‐based framework synthesizing models across these two fields is lacking. Here we review 65 representative community and invasion models and propose a common framework articulated around six processes (dispersal, drift, abiotic interactions, within‐guild interactions, cross‐guild interactions, and genetic changes). The framework is designed to synthesize the content of the two fields, provide a general perspective on their development, and enable their comparison. The application of this framework and of a novel method based on network theory reveals some lack of coherence between the two fields, despite some historical similarities. Community ecology models are characterized by combinations of multiple processes, likely reflecting the search for an overarching theory to explain community assembly and structure, drawing predominantly on interaction processes, but also accounting largely for the other processes. In contrast, most models in invasion ecology invoke fewer processes and focus more on interactions between introduced species and their novel biotic and abiotic environment. The historical dominance of interaction processes and their independent developments in the two fields is also reflected in the lower level of coherence for models involving interactions, compared to models involving dispersal, drift, and genetic changes. It appears that community ecology, with a longer history than invasion ecology, has transitioned from the search for single explanations for patterns observed in nature to investigate how processes may interact mechanistically, thereby generating and testing hypotheses. Our framework paves the way for a similar transition in invasion ecology, to better capture the dynamics of multiple alien species introduced in complex communities. Reciprocally, applying insights from invasion to community ecology will help us understand and predict the future of ecological communities in the Anthropocene, in which human activities are weakening species’ natural boundaries. Ultimately, the successful integration of the two fields could advance a predictive ecology that is urgently required in a rapidly changing world.

The Table Mountain Ghost Frog (Heleophryne rosei) is endemic to the Table Mountain massif and is Critically Endangered. Other than clear, clean perennial stream flow, the optimal aquatic conditions required by their larvae are unknown. Dissolved oxygen, temperature, pH, electro-conductivity, aspect and permanence of flow are the independent variables measured seasonally at two sampling altitudes at 12 rivers of the massif. Using a logistic regression model we found that a permanence of water flow and lower water temperature were significant predictors of tadpole presence. Streams with mean summer temperature above 17.2 °C, at 300 m – 400 m above sea level, do not have tadpoles. Summer and autumn abstraction should be avoided, while a summer water temperature above an average of 17.2 °C is a threshold of potential concern for management authorities responsible for biodiversity conservation, threat mitigation efforts, and bulk-water supply and abstraction. Conservation implications: The Environmental Water Reserve has not been determined for streams of Table Mountain. The requirements of the Critically Endangered Table Mountain Ghost Frog (Heleophryne rosei) can be adopted as the minimum conditions to support this species and associated communities. Perennial flow, an average January water temperature of 17.2 °C or lower.

It is essential to incorporate mechanisms of biotic resistance into predictions of ecological impact conferred by invasive species. Trophically and functionally analogous native species have high potential to confer biotic resistance or to be susceptible to competitive exclusion by the invading species. In species with dominance hierarchies and high aggression the role of weapons, such as chelae, is emphasised and selected for. Differences in traits such as crushing capacity can indicate prey handling capabilities, while correlations between closing force and morphology can be used to understand the role of signaling in agonistic contests. Closing force strength can be used to infer the outcomes of both direct (predation) and indirect (competition) trophic interactions. Southern Africa has been invaded by two freshwater crayfish species (Cherax quadricarinatus and Procambarus clarkii). Biotic resistance of freshwater crabs toward crayfish invasions varies between geographic location and co-evolutionary history, thus comparing invasion histories without incorporating geographic context can produce unequivocal conclusions. We compared the closing force and chelae morphology of both crayfish species with a native trophically analogous freshwater crab, Potamonautes perlatus. Closing force increased significantly with mass for all species. There was significant interaction between sex and species on closing force. Potamonautes perlatus females showed significantly stronger maximum chela closing force than male P. perlatus, both sexes of P. clarkii and female C. quadricarinatus. Contrastingly, male C. quadricarinatus had significantly higher closing forces than both sexes of P. clarkii and female C. quadricarinatus, however, there was no difference between female P. perlatus. Native P. perlatus has the capacity to hold a competitive mechanical advantage over both invaders, but this varies with sex. Chelae length was not a significant predictor for closing strength in any of the species, which may be related to dishonest signaling in decapod species. This makes it imperative to assess whether factors such as closing force actually translate to resource holding potential in a contest scenario. We thus provide evidence that African freshwater crabs may exhibit biotic resistance toward invasion and the first measurements for C. quadricarinatus closing force strength.

Invasive alien species (IAS) are a growing threat globally and cause a variety of ecological, economic, and social impacts. People can introduce IAS and facilitate their spread, and can also implement, support, or oppose their management. Understanding local knowledge, awareness, and perceptions are therefore crucial if management and policy are to be effective. We administered questionnaires to members of the public in eight small towns along the Berg River Catchment in the biodiverse fynbos biome of South Africa. We aimed to assess: (1) awareness of IAS by the general public, (2) local perceptions of the impacts associated with IAS, (3) whether awareness of IAS is correlated with demographic covariates and IAS density, and (4) people’s willingness to detect, report, and support IAS management. Overall, 262 respondents participated in the survey. Most respondents (65%) did not know what IAS are, and 10% were unsure. Many respondents also perceived IAS as beneficial. Using a logistic regression, we found that IAS density, educational level, and gender influenced people’s knowledge and perceptions about IAS in the region. There were a small number (4%) of respondents currently detecting and reporting IAS, but many respondents were interested to learn more. We concluded that people living in small towns in the Western Cape of South Africa remain largely unaware of IAS and their impacts. It is crucial to increase awareness-raising initiatives, and build support and engagement in management of IAS in small towns.

Temperature and water availability are two of the most important variables affecting all aspects of an anuran’s key physiological processes such as body temperature (Tb), evaporative water loss (EWL) and standard metabolic rate (SMR). Since anurans display pronounced sexual dimorphism, evidence suggests that these processes are further influenced by other factors such as vapour pressure deficit (VPD), sex and body mass (Mb). However, a limited number of studies have tested the generality of these results across a wide range of ecologically relevant ambient temperatures (Ta), while taking habitat use into account. Thus, the aim of this study was to investigate the role of Ta on Tb, whole-animal EWL and whole-animal SMR in three wild caught African anuran species with different ecological specialisations: the principally aquatic African clawed frog (Xenopus laevis), stream-breeding common river frog (Amietia delalandii), and the largely terrestrial raucous toad (Sclerophrys capensis). Experiments were conducted at a range of test temperatures (5–35 °C, at 5 °C increments). We found that VPD better predicted rates of EWL than Ta in two of the three species considered. Moreover, we found that Tb, whole-animal EWL and whole-animal SMR increased with increasing Ta, while Tb increased with increasing Mb in A. delalandii and S. capensis but not in X. laevis. Whole-animal SMR increased with increasing Mb in S. capensis only. We did not find any significant effect of VPD, Mb or sex on whole-animal EWL within species. Lastly, Mb did not influence Tb, whole-animal SMR and EWL in the principally aquatic X. laevis. These results suggest that Mb may not have the same effect on key physiological variables, and that the influence of Mb may also depend on the species ecological specialisation. Thus, the generality of Mb as an important factor should be taken in the context of both physiology and species habitat specialisation.

Developed countries are producing policies to reduce the flow of invasive species and control or eradicate existing invasions, but most developing countries are under-resourced to tackle either aspect without help. Emerging economies, such as Brazil, Russia, India, China, and South Africa (BRICS), are responsible for donating many of the world's invasive species that have the potential to reach nearly all terrestrial biomes. Implementing a proactive 'facilitated network' model is urgently required to build capacity and stimulate effective appropriate invasion science. We contend that creating a BRICS network of invasion scientists will have the immediate impact required to meet future policy demands.

Background: Conservation relies on the strategic use of resources because monies for conservation action are limited, especially in developing countries. South Africa’s Frog Atlas project established a baseline for the country’s amphibian data and threat levels in 2004, and in 2009 a prioritisation exercise developed a strategy for conservation research. Objectives: In this article, we assess this strategy for conservation research. Method: We conducted a quantitative and qualitative assessment of research undertaken since the strategy was developed. Results: The strategy has produced a lasting impact on taxonomy, ecological studies, monitoring and capacity building. Publications in all areas have increased, but particularly in conservation ecology. Other indicators are increases in the numbers of locality records for target taxa, species descriptions and postgraduate degrees with amphibians as the principal topic. We document important milestones for South African amphibian conservation, including the first Biodiversity Management Plan for Species (BMP-S) for Hyperolius pickersgilli, a smart device app that uploads locality data to an open access database, 15 years of monitoring data and new amphibian identification books for adults and children. The Red List Index calculated for South African amphibians shows that the country’s species are becoming more threatened (a 1% reduction in 10 years), but a hindcasting exercise suggests that most of the damage was already done by 1990. We provide a checklist for 131 amphibian species in South Africa, of which 82 species are endemic. Conclusion: A strategy for conservation research was found to greatly augment the focus of research on South African frogs. A new strategy should focus on fewer taxa over meaningful time spans.

The study and management of emerging infectious diseases (EIDs) and of biological invasions both address the ecology of human-associated biological phenomena in a rapidly changing world. However, the two fields work mostly in parallel rather than in concert. This review explores how the general phenomenon of an organism rapidly increasing in range or abundance is caused, highlights the similarities and differences between research on EIDs and invasions, and discusses shared management insights and approaches. EIDs can arise by: (i) crossing geographical barriers due to human-mediated dispersal, (ii) crossing compatibility barriers due to evolution, and (iii) lifting of environmental barriers due to environmental change. All these processes can be implicated in biological invasions, but only the first defines them. Research on EIDs is embedded within the One Health concept—the notion that human, animal and ecosystem health are interrelated and that holistic approaches encompassing all three components are needed to respond to threats to human well-being. We argue that for sustainable development, biological invasions should be explicitly considered within One Health. Management goals for the fields are the same, and direct collaborations between invasion scientists, disease ecologists and epidemiologists on modelling, risk assessment, monitoring and management would be mutually beneficial.

The desire to own a pet amphibian is growing, and with it a growth in amphibian trade and in negative impacts on native populations, including disease transmission and invasive amphibian populations. We know very little about how or why people choose amphibians as pets, but amphibian owners share large numbers of videos on freely accessible platforms, such as YouTube. We aimed to use videos of captive amphibians to determine which species are kept, their life-history stage and the types of videos uploaded. We watched and categorized 1,162 videos by video type, type of amphibian behavior and amphibian taxonomy (superfamily, family, and species). We used data on the amphibian trade from the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), on conservation status from the International Union for Conservation of Nature (IUCN) red list, and on potential environmental impact from published Environmental Impact Classification of Alien Taxa (EICAT) records, to determine potential conflicts of owning pet amphibians. We recorded 173 captive species in 847 videos with a taxonomic overrepresentation of salamandroids and pipoids, and an underrepresentation of ranoids and plethodontoids. When compared to videos of wild amphibian species, videos of captive animals featured disproportionate amounts of adults feeding, being handled and moving. The videos watched had a smaller proportion of threatened amphibian species, but a higher proportion of invasive species, than would be expected by chance, with the proportion present in CITES appendices (18%) being non-significant. We suggest that such data can be used to profile potential pets for trade and attempt to avoid conflicts with threatened and highly impacting alien species.

For many taxa, new records of non-native introductions globally occur at a near exponential rate. We undertook a systematic review of peer-reviewed publications on non-native herpetofauna, to assess the information base available for assessing risks of future invasions, resulting in 836 relevant papers. The taxonomic and geographic scope of the literature was also compared to a published database of all known invasions globally. We found 1,116 species of herpetofauna, 95% of which were present in fewer than 12 studies. Nearly all literature on the invasion ecology of herpetofauna has appeared since 2000, with a strong focus on frogs (58%), particularly cane toads (Rhinella marina) and their impacts in Australia. While fewer papers have been published on turtles and snakes, proportionately more species from both these groups have been studied than for frogs. Within each herpetofaunal group, there are a handful of well-studied species: R. marina, Lithobates catesbeianus, Xenopus laevis, Trachemys scripta, Boiga irregularis and Anolis sagrei. Most research (416 papers; 50%) has addressed impacts, with far fewer studies on aspects like trade (2%). Besides Australia (213 studies), most countries have little location-specific peer-reviewed literature on non-native herpetofauna (on average 1.1 papers per established species). Other exceptions were Guam, the UK, China, California and France, but even their publication coverage across established species was not even. New methods for assessing and prioritizing invasive species such as the Environmental Impact Classification for Alien Taxa provide useful frameworks for risk assessment, but require robust species-level studies. Global initiatives, similar to the Global Amphibian Assessment, using the species and taxonomic groups identified here, are needed to derive the level of information across broad geographic ranges required to apply these frameworks. Expansive studies on model species can be used to indicate productive research foci for understudied taxa.

Predators can play an important role in regulating prey abundance and diversity, determining food web structure and function, and contributing to important ecosystem services, including the regulation of agricultural pests and disease vectors. Thus, the ability to predict predator impact on prey is an important goal in ecology. Often, predators of the same species are assumed to be functionally equivalent, despite considerable individual variation in predator traits known to be important for shaping predator–prey interactions, like body size. This assumption may greatly oversimplify our understanding of within-species functional diversity and undermine our ability to predict predator effects on prey. Here, we examine the degree to which predator–prey interactions are functionally homogenous across a natural range of predator body sizes. Specifically, we quantify the size-dependence of the functional response of African clawed frogs (Xenopus laevis) preying on mosquito larvae (Culex pipiens). Three size classes of predators, small (15–30 mm snout-vent length), medium (50–60 mm) and large (105–120 mm), were presented with five densities of prey to determine functional response type and to estimate search efficiency and handling time parameters generated from the models. The results of mesocosm experiments showed that type of functional response of X. laevis changed with size: small predators exhibited a Type II response, while medium and large predators exhibited Type III responses. Functional response data showed an inversely proportional relationship between predator attack rate and predator size. Small and medium predators had highest and lowest handling time, respectively. The change in functional response with the size of predator suggests that predators with overlapping cohorts may have a dynamic impact on prey populations. Therefore, predicting the functional response of a single size-matched predator in an experiment may misrepresent the predator’s potential impact on a prey population.

Amphibian invasions have considerable detrimental impacts on recipient ecosystems. However, reliable risk analysis of invasive amphibians still requires research on more non-native amphibian species. An invasive population of the Indian bullfrog, Hoplobatrachus tigerinus, is currently spreading on the Andaman archipelago and may have significant trophic impacts on native anurans through competition and predation. We carried out diet analyses of the invasive H. tigerinus and native anurans, across four habitat types and two seasons; we hypothesized that (i) small vertebrates constitute a majority of the H. tigerinus diet, particularly by volume and (ii) the diet of H. tigerinus significantly overlaps with the diet of native anurans, thereby, leading to potential competition. We assessed the diet of the invasive H. tigerinus (n = 358), and individuals of the genera Limnonectes (n = 375) and Fejervarya (n = 65) and found a significant dietary overlap of H. tigerinus with only Limnonectes. Small vertebrates, including several endemic species, constituted the majority of H. tigerinus, diet by volume, suggesting potential impact by predation. Prey consumption and electivity of the three anurans indicated a positive relationship between predator-prey body sizes. Individuals of H. tigerinus and Fejervarya chose evasive prey, suggesting that these two taxa are mostly ambush predators; individuals of Limnonectes chose a mixture of sedentary and evasive prey indicating that the species employs a combination of ‘active search’ and ‘sit and wait’ foraging strategies. All three species of anurans mostly consumed terrestrial prey. This intensive study on a genus of newly invasive amphibian contributes to knowledge of the impact of amphibian invasions, and elucidates the feeding ecology of H. tigerinus, and species of the genera Limnonectes and Fejervarya. We also stress the necessity to evaluate prey availability and volume in future studies for meaningful insights into diet of amphibians.

Dispersal forms are an important component of the ecology of many animals, and reach particular importance for predicting ranges of invasive species. African clawed frogs (Xenopus laevis) move overland between water bodies, but all empirical studies are from invasive populations with none from their native southern Africa. Here we report on incidents of overland movement found through a capture-recapture study carried out over a three year period in Overstrand, South Africa. The maximum distance moved was 2.4 km with most of the 91 animals, representing 5% of the population, moving ∼150 m. We found no differences in distances moved by males and females, despite the former being smaller. Fewer males moved overland, but this was no different from the sex bias found in the population. In laboratory performance trials, we found that males outperformed females, in both distance moved and time to exhaustion, when corrected for size. Overland movement occurred throughout the year, but reached peaks in spring and early summer when temporary water bodies were drying. Despite permanent impoundments being located within the study area, we found no evidence for migrations of animals between temporary and permanent water bodies. Our study provides the first dispersal kernel for X. laevis and suggests that it is similar to many non-pipid anurans with respect to dispersal.

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.

The widespread African clawed frog (Xenopus laevis) occurs in sympatry with the IUCN Endangered Cape platanna (Xenopus gilli) throughout its entire range in the south-western Cape, South Africa. In order to investigate aspects of the interspecific competition between populations of X. laevis and X. gilli, an assessment of their niche differentiation was conducted through a comprehensive study on food composition and trophic niche structure at two study sites: the Cape of Good Hope (CoGH) and Kleinmond. A total of 399 stomach contents of X. laevis (n = 183) and X. gilli (n = 216) were obtained together with samples of available prey to determine food preferences using the Electivity index (E*), the Simpson’s index of diversity (1 − D), the Shannon index (H′), and the Pianka index (Ojk). Xenopus gilli diet was more diverse than X. laevis, particularly in Kleimond where the Shannon index was nearly double. Both species were found to consume large amounts of tadpoles belonging to different amphibian species, including congeners, with an overall higher incidence of anurophagy than previously recorded. However, X. laevis also feeds on adult X. gilli, thus representing a direct threat for the latter. While trophic niche overlap was 0.5 for the CoGH, it was almost 1 in Kleinmond, suggesting both species utilise highly congruent trophic niches. Further, subdividing the dataset into three size classes revealed overlap to be higher in small frogs in both study sites. Our study underlines the importance of actively controlling X. laevis at sites with X. gilli in order to limit competition and predation, which is vital for conservation of the south-western Cape endemic.

Background: Globally, invasive amphibians are known for their environmental and social impacts that range from poisoning of local fauna and human populations to direct predation on other amphibians. Although several countries on most continents have had multiple introductions of many species, southern Africa appears to have escaped allochthonous introductions. Instead, it has a small number of domestic exotic species that have rapidly expanded their ranges and established invasive populations within South Africa. Objectives & methods: We used the literature to provide a historical overview of dispersal by some of the world’s major invasive amphibians, give examples of species that are commonly moved as stowaways and discuss historical and current amphibian trade in the region. In addition, we give an overview of new South African legislation and how this is applied to amphibian invasions, as well as providing updates on the introduced populations of three domestic exotics: Hyperolius marmoratus, Sclerophrys gutturalis and Xenopus laevis. Results: We show that frogs are mainly moved around southern Africa through ‘jump’ dispersal, although there are a number of records of ‘cultivation’, ‘leading-edge’ and ‘extreme long-distance’ dispersal types. Important pathways include trade in fruit and vegetables, horticultural products and shipping containers. Conclusion: We suggest that southern Africa is becoming more vulnerable to amphibian invasions because of an increase in trade, agricultural and domestic impoundments as well as global climate change. Increasing propagule pressure suggests that preventing new introductions will become a key challenge for the future. Currently, trade in amphibians in the region is practically non-existent, suggesting potential for best practice to prevent importation of species with high invasion potential and to stop the spread of disease.

African clawed frogs (Xenopus laevis) are often referred to as ‘purely aquatic’ but there are many publications which suggest extensive overland movements. Previous reviews which considered the topic have not answered the following questions: (1) is there evidence for overland dispersal in native and invasive ranges; (2) what is the range of distances moved overland; (3) when does overland movement occur; and (4) is there evidence of breeding migratory behaviour? A systematic review was chosen to synthesise and critically analyse all literature on the overland movement in Xenopus laevis. Database searches resulted in 57 documents which revealed a paucity of empirical studies, with 28 containing no data, and 19 having anecdotal content. Overwhelming evidence shows that both native and invasive populations of X. laevis move overland, with well documented examples for several other members of the genus (X. borealis, X. gilli, X. muelleri, X. fraseriand X. tropicalis). Reports of distances moved overland were from 40 m to 2 km, with no apparent difference between native and invasive ranges. Overland movements are not confined to wet seasons or conditions, but the literature suggests that moving overland does not occur in the middle of the day. Migrations to temporary water-bodies for breeding have been suggested, but without any corroborating data.

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Many animals show unique morphological and behavioural adaptations to specific habitats. In particular, variation in cranial morphology is known to influence feeding performance, which in turn influences dietary habits and, ultimately, fitness. Dietary separation is an important means of partitioning ecological niches and avoiding inter- and intraspecific competition. Consequently, differences in dietary resources may help explain phenotypic divergence in closely-related species occupying different habitats, as well as sexual dimorphism. We test this hypothesis on five phenotypic forms of a recent radiation of dwarf chameleons ( Bradypodion) that vary extensively in habitat use and cranial morphology. By examining stomach contents, the dietary composition of each phenotypic form is compared to investigate potential differences in feeding strategies. Overall, chameleons in the present study exhibit considerable dietary overlap (at both inter- and intraspecific levels), indicating that diet is not a major driver of variation in cranial morphology within this radiation. However, the stomachs of closed-canopy females were found to contain more prey items than male stomachs, possibly indicating that females require a greater caloric intake than their male counterparts. [ABSTRACT FROM AUTHOR]

Two generations of gustatory organs occur during amphibian ontogeny in frogs and salamanders (Anura and Caudata), and are classified as taste buds or taste discs. Taste buds are present in larval forms, whereas taste discs are typical for adults. The little research done on Gymnophiona suggests that only taste buds are present in aquatic forms (larvae and adults), and adults of the only terrestrial species studied had taste buds. Here we investigate the nature of gustatory organs in a terrestrial caecilian, Boulengerula boulengeri (Herpelidae), from Tanzania using standard light and scanning electron microscopy. We found only taste bud type organs to be present in B. boulengeri. These occur mainly in the mucosa of the front part of the oral cavity, usually near the teeth. Our results suggest that B. boulengeri possesses only one type of gustatory organ during its ontogeny. [ABSTRACT FROM AUTHOR]

Convergent evolution can explain similarity in morphology between species, due to selection on a fitness-enhancing phenotype in response to local environmental conditions. As selective pressures on body morphology may be strong, these have confounded our understanding of the evolutionary relationships between species. Within the speciose African radiation of lacertid lizards (Eremiadini), some species occupy a narrow habitat range (e.g. open habitat, cluttered habitat, strictly rupicolous, or strictly psammophilic), which may exert strong selective pressures on lizard body morphology. Here we show that the overall body plan is unrelated to shared ancestry in the African radiation of Eremiadini, but is instead coupled to habitat use. Comprehensive Bayesian and likelihood phylogenies using multiple representatives from all genera (2 nuclear, 2 mitochondrial markers) show that morphologically convergent species thought to represent sister taxa within the same genus are distantly related evolutionary lineages (Ichnotropis squamulosa and Ichnotropis spp.; Australolacerta rupicola and A. australis). Hierarchical clustering and multivariate analysis of morphological characters suggest that body, and head, width and height (stockiness), all of which are ecologically relevant with respect to movement through habitat, are similar between the genetically distant species. Our data show that convergence in morphology, due to adaptation to similar environments, has confounded the assignment of species leading to misidentification of the taxonomic position of I. squamulosa and the Australolacerta species. [ABSTRACT FROM AUTHOR]

Aim Amphibians are a model group for studies of the biogeographical origins of salt-intolerant taxa on oceanic islands. We used the Gulf of Guinea islands to explore the biogeographical origins of island endemism of one species of frog, and used this to gain insights into potential colonization mechanisms. Location São Tomé and Príncipe, two of the four major islands in the Gulf of Guinea, West Africa, are truly oceanic and have an exceptionally high biodiversity. Methods Mitochondrial DNA is used to test the endemic status of a frog from São Tomé and compare it with congeneric taxa from tropical Africa. Existing data on surface currents, surface salinity, atmospheric circulation and bird migration in the Gulf of Guinea are summarized to address hypotheses concerning colonization mechanisms. Results The endemic status of Ptychadena newtoni (Bocage) is supported here by mitochondrial DNA sequences, and analysis of this and other molecular data indicates that an East African species close to Ptychadena mascareniensis (Duméril and Bibron) is its nearest relative. We refute the possibility that this population was anthropogenically introduced, in favour of a natural dispersal mechanism. Main conclusions With six endemic frogs and one caecilian, the Gulf of Guinea islands harbour a diverse amphibian fauna. Five of these species appear to have their closest relatives in East Africa. Insufficient evidence exists for transportation by storms, birds or rafts alone. However, we propose a synergy of rafting, favourable surface currents and a reduction in salinity of surface waters. Catastrophic events, or wet periods in climatic history, could allow freshwater paths to open far enough to enable continental flora and fauna to reach these and other isolated oceanic islands. [ABSTRACT FROM AUTHOR]

Boulengerula uluguruensis is a terrestrial caeciliid caecilian described from the Uluguru Mountains of Tanzania. We investigated the relative abundance of B. uluguruensis in agricultural and forested habitats at the beginning of the rainy season. This caecilian was found over a wide altitudinal range (450–1175 m a.s.l.), and in many land cover types (including natural forest, plantation forest and small scale agriculture), and different soil textures (including very compact soil). Based on quantitative and semi-quantitative surveys, B. uluguruensis is more abundant than any other subterranean lower vertebrate in this area, with densities up to 0.4 individuals m−2 in some agricultural plots. The hypothesis proposed, that B. uluguruensis is more abundant in agriculture than their native forest, could not be conclusively tested during this brief visit, although the data that were collected do not indicate to the contrary. Likewise, the size of animals from forest and agricultural populations could not be objectively compared. [ABSTRACT FROM AUTHOR]

The potentially important ecology of subterranean predators of soil ecosystem engineers is poorly understood. This is especially true of caecilian amphibians (Gymnophiona) for which there are virtually no quantitative data. Results of the first field trials of permanent marking in caecilians are presented. A preliminary assessment is made of the efficacy of mark–recapture studies for estimating population size of Gegeneophis ramaswamii Taylor in 100 m2 of low intensity agriculture in Kerala, India. Over three sampling occasions spanning 58 days of the monsoon season, 114 individuals were captured, 104 marked and released, and 21 recaptured. Models estimate an open population of 60 individuals (95% confidence interval of 45.2 to 151.3), and a closed population of 236 (95% confidence interval of 174 to 351). A census interpretation of the raw capture data gives densities of about 0.31 to 0.48 m-2. Results suggest large movement in and out of the sampled area during the study. Despite caveats associated with these data, progress is made in identifying potential limitations and improvements in the methods used. This study highlights the paucity of knowledge of caecilian ecology, and the need for long-term studies to elucidate further ecological information and to monitor populations. [ABSTRACT FROM AUTHOR]

Climatic changes have had profound impacts on species distributions throughout time. In response, species have shifted ranges, adapted genetically and behaviourally or become extinct. Using species distribution models, we examined how changes in suitable climatic space could affect the distributions of 37 endemic frog species in the Cape Floristic Region (CFR) - an area proposed to have evolved its megadiversity under a stable climate, which is expected to change substantially in future. Species distributions were projected onto mean climate for a current period (1950 to 2000), hindcasted to palaeoclimate (Last Glacial Maximum; LGM ≈ 21 kya and Holocene Glacial Minimum; HGM ≈ 6 kya) and forecasted for two emissions scenarios (A2a and B2a) for the year 2080. We then determined the changes in area sizes, direction (longitude and latitude), fragmentation index and biotic velocity, and assessed if these were affected by life-history traits and altitude. We found that the biotic velocity at which the CFR amphibian community is expected to shift north (A2a ≈ 540.5 km/kya) and east (B2a ≈ 198 km/kya) far exceeds historical background rates (≈1.05 km/kya, north and west ≈ 2.36 km/kya since the LGM). Our models further suggest that the CFR amphibian community has already lost about 56% of suitable climate space since the LGM and this loss is expected to accelerate under future emission scenarios (A2a ≈ 70%; B2a ≈ 60%). Lastly, we found that highland species were more fragmented than lowland species between the LGM and current period, but that the fragmentation of lowland species between current and future climates is expected to increase. [ABSTRACT FROM AUTHOR]

Abstract The Cape platanna, Xenopus gilli, an endangered frog, hybridizes with the African clawed frog, X. laevis, in South Africa. Estimates of the extent of gene flow between these species range from pervasive to rare. Efforts have been made in the last 30 years to minimize hybridization between these two species in the west population of X. gilli, but not the east populations. To further explore the impact of hybridization and the efforts to minimize it, we examined molecular variation in one mitochondrial and 13 nuclear genes in genetic samples collected recently (2013) and also over two decades ago (1994). Despite the presence of F 1 hybrids, none of the genomic regions we surveyed had evidence of gene flow between these species, indicating a lack of extensive introgression. Additionally we found no significant effect of sampling time on genetic diversity of populations of each species. Thus, we speculate that F 1 hybrids have low fitness and are not backcrossing with the parental species to an appreciable degree. Within X. gilli, evidence for gene flow was recovered between eastern and western populations, a finding that has implications for conservation management of this species and its threatened habitat.

Classification of alien species’ impacts can aid policy making through evidence based listing and management recommendations. We highlight differences and a number of potential difficulties with two scoring tools, the Environmental Impact Classification of Alien Taxa (EICAT) and the Generic Impact Scoring System (GISS) using amphibians as a case study. Generally, GISS and EICAT assessments lead to very similar impact levels, but scores from the schemes are not equivalent. Small differences are attributable to discrepancies in the verbal descriptions for scores. Differences were found in several impact categories. While the issue of disease appears to be related to uncertainties in both schemes, hybridisation might be inflated in EICAT. We conclude that GISS scores cannot directly be translated into EICAT classifications, but they give very similar outcomes and the same literature base can be used for both schemes.

Background The Australian redclaw crayfish (Cherax quadricarinatus, von Martens), is native to Australasia, but has been widely translocated around the world due to aquaculture and aquarium trade. Mostly as a result of escape from aquaculture facilities, this species has established extralimital populations in Australia and alien populations in Europe, Asia, Central America and Africa. In South Africa, C. quadricarinatus was first sampled from the wild in 2002 in the Komati River, following its escape from an aquaculture facility in Swaziland, but data on the current status of its populations are not available. Methods To establish a better understanding of its distribution, rate of spread and population status, we surveyed a total of 46 sites in various river systems in South Africa and Swaziland. Surveys were performed between September 2015 and August 2016 and involved visual observations and the use of collapsible crayfish traps. Results Cherax quadricarinatus is now present in the Komati, Lomati, Mbuluzi, Mlawula and Usutu rivers, and it was also detected in several off-channel irrigation impoundments. Where present, it was generally abundant, with populations having multiple size cohorts and containing ovigerous females. In the Komati River, it has spread more than 112 km downstream of the initial introduction point and 33 km upstream of a tributary, resulting in a mean spread rate of 8 km year−1 downstream and 4.7 km year−1 upstream. In Swaziland, estimated downstream spread rate might reach 14.6 km year−1. Individuals were generally larger and heavier closer to the introduction site, which might be linked to juvenile dispersal. Discussion These findings demonstrate that C. quadricarinatus is established in South Africa and Swaziland and that the species has spread, not only within the river where it was first introduced, but also between rivers. Considering the strong impacts that alien crayfish usually have on invaded ecosystems, assessments of its potential impacts on native freshwater biota and an evaluation of possible control measures are, therefore, urgent requirements.

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.

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.

Background. Frogs are generalist predators of a wide range of typically small prey items. But descriptions of dietary items regularly include other anurans, such that frogs are considered to be among the most important of anuran predators. However, the only existing hypothesis for the inclusion of anurans in the diet of post-metamorphic frogs postulates that it happens more often in bigger frogs. Moreover, this hypothesis has yet to be tested.Methods. We reviewed the literature on frog diet in order to test the size hypothesis and determine whether there are other putative explanations for anurans in the diet of post-metamorphic frogs. In addition to size, we recorded the habitat, the number of other sympatric anuran species, and whether or not the population was invasive. We controlled for taxonomic bias by including the superfamily in our analysis.Results. Around one fifth of the 355 records included anurans as dietary items of populations studied, suggesting that frogs eating anurans is not unusual. Our data showed a clear taxonomic bias with ranids and pipids having a higher proportion of anuran prey than other superfamilies. Accounting for this taxonomic bias, we found that size in addition to being invasive, local anuran diversity, and habitat produced a model that best fitted our data. Large invasive frogs that live in forests with high anuran diversity are most likely to have a higher proportion of anurans in their diet.Conclusions. We confirm the validity of the size hypothesis for anurophagy, but show that there are additional significant variables. The circumstances under which frogs eat frogs are likely to be complex, but our data may help to alert conservationists to the possible dangers of invading frogs entering areas with threatened anuran species.

Islands provide exciting opportunities for exploring ecological and evolutionary mechanisms. The oceanic island of São Tomé in the Gulf of Guinea exhibits high diversity of fauna including the endemic caecilian amphibian, Schistometopum thomense. Variation in pigmentation, morphology and size of this taxon over its c. 45 km island range is extreme, motivating a number of taxonomic, ecological, and evolutionary hypotheses to explain the observed diversity. We conducted a population genetic study of S. thomense using partial sequences of two mitochondrial DNA genes (ND4 and 16S), together with morphological examination, to address competing hypotheses of taxonomic or clinal variation. Using Bayesian phylogenetic analysis and Spatial Analysis of Molecular Variance, we found evidence of four geographic clades, whose range and approximated age (c. 253 Kya-27 Kya) are consistent with the spread and age of recent volcanic flows. These clades explained 90% of variation in ND4 (φCT = 0.892), and diverged by 4.3% minimum pairwise distance at the deepest node. Most notably, using Mismatch Distributions and Mantel Tests, we identified a zone of population admixture that dissected the island. In the northern clade, we found evidence of recent population expansion (Fu's Fs = -13.08 and Tajima's D = -1.80) and limited dispersal (Mantel correlation coefficient = 0.36, p = 0.01). Color assignment to clades was not absolute. Paired with multinomial regression of chromatic data, our analyses suggested that the genetic groups and a latitudinal gradient together describe variation in color of S. thomense. We propose that volcanism and limited dispersal ability are the likely proximal causes of the observed genetic structure. This is the first population genetic study of any caecilian and demonstrates that these animals have deep genetic divisions over very small areas in accordance with previous speculations of low dispersal abilities.

Phenotypic performance in different environments is central to understanding the evolutionary and ecological processes that drive adaptive divergence and, ultimately, speciation. Because habitat structure can affect an animal's foraging behaviour, anti-predator defences, and communication behaviour, it can influence both natural and sexual selection pressures. These selective pressures, in turn, act upon morphological traits to maximize an animal's performance. For performance traits involved in both social and ecological activities, such as bite force, natural and sexual selection often interact in complex ways, providing an opportunity to understand the adaptive significance of morphological variation with respect to habitat. Dwarf chameleons within the Bradypodion melanocephalum-Bradypodion thamnobates species complex have multiple phenotypic forms, each with a specific head morphology that could reflect its use of either open- or closed-canopy habitats. To determine whether these morphological differences represent adaptations to their habitats, we tested for differences in both absolute and relative bite performance. Only absolute differences were found between forms, with the closed-canopy forms biting harder than their open-canopy counterparts. In contrast, sexual dimorphism was found for both absolute and relative bite force, but the relative differences were limited to the closed-canopy forms. These results indicate that both natural and sexual selection are acting within both habitat types, but to varying degrees. Sexual selection seems to be the predominant force within the closed-canopy habitats, which are more protected from aerial predators, enabling chameleons to invest more in ornamentation for communication. In contrast, natural selection is likely to be the predominant force in the open-canopy habitats, inhibiting the development of conspicuous secondary sexual characteristics and, ultimately, enforcing their overall diminutive body size and constraining performance.

Low-lying areas of the Cape at Africa’s south-westernmost tip have undergone dramatic marine-remodelling, with regular changes in sea-level following glacial cycles. Species for which marine barriers are impenetrable underwent concomitant radical distribution changes which may account for current range disjunctions. The Cape platanna, Xenopus gilli, is a frog distributed in only three disjunt areas within low-lying regions of the southwestern Cape. We determined the relationship between frogs from these three disjunct areas, by using a combination of morphometric analysis and mtDNA (ND2 and 16S fragments) sequences of 130 frogs from eight ponds. Coalescent analyses on molecular data dated the divergence in two major clades to around 4.6 Mya, a period during which major uplifting on the eastern side of the subcontinent caused climate changes throughout southern Africa. Principal components analysis showed significant morphometric differences between each clade on head and limb measurements. Consistent differences in ventral colouration and patterning were also observed. We report on increased levels of hybridisation with X. laevis throughout the range of X. gilli, which reaches at least 27% hybrids in some ponds. Urgent conservation actions are required to control habitat loss from alien invasive vegetation, and prevent introgression with the domestic-exotic, X. laevis.

No direct palaeoclimatic proxies have been available to indicate the seasonality or amount of rainfall on the west coast of southern Africa during the Early Pliocene. The Benguela Upwelling System (BUS) is today one of the factors responsible for the present-day aridity on the west coast of southern Africa. The initiation of the BUS is frequently linked to the entrenchment of aridity and the establishment of the current winter rainfall pattern on the west coast; however, marine proxies are inconclusive regarding the effects of past fluctuations in the BUS and sea surface temperatures on the rainfall regime. Neither the fossil evidence nor the fact that plants using the C3 photosynthetic pathway predominate at this time, provide direct evidence of winter rainfall at Langebaanweg. We challenge certain assumptions which are commonly made in the literature regarding the timing of inception of a winter rainfall regime on the west coast and the onset of aridity in the Langebaan region, and provide new evidence as to seasonality of rainfall at Langebaanweg in the Early Pliocene. Herein, the identification of frog species from the genus Ptychadena from Langebaanweg provides new and compelling evidence for a summer rainfall regime, or of at least significant summer rainfall, at 5.1 mya in the southwestern Cape of South Africa. Significance: • Advances understanding of the evolution of the winter rainfall zone on the west coast of South Africa • Assesses evidence for the inception of aridity and a winter rainfall regime on the west coast of South Africa • Fossil Ptychadenidae from the Early Pliocene site of Langebaanweg provide evidence for a summer rainfall regime at 5.1 mya on the west coast of the southwestern Cape.

Convergent evolution can explain similarity in morphology between species, due to selection on a fitness-enhancing phenotype in response to local environmental conditions. As selective pressures on body morphology may be strong, these have confounded our understanding of the evolutionary relationships between species. Within the speciose African radiation of lacertid lizards (Eremiadini), some species occupy a narrow habitat range (e.g. open habitat, cluttered habitat, strictly rupicolous, or strictly psammophilic), which may exert strong selective pressures on lizard body morphology. Here we show that the overall body plan is unrelated to shared ancestry in the African radiation of Eremiadini, but is instead coupled to habitat use. Comprehensive Bayesian and likelihood phylogenies using multiple representatives from all genera (2 nuclear, 2 mitochondrial markers) show that morphologically convergent species thought to represent sister taxa within the same genus are distantly related evolutionary lineages (Ichnotropis squamulosa and Ichnotropis spp.; Australolacerta rupicola and A. australis). Hierarchical clustering and multivariate analysis of morphological characters suggest that body, and head, width and height (stockiness), all of which are ecologically relevant with respect to movement through habitat, are similar between the genetically distant species. Our data show that convergence in morphology, due to adaptation to similar environments, has confounded the assignment of species leading to misidentification of the taxonomic position of I. squamulosa and the Australolacerta species.