Functional responses can’t unify invasion ecology

Dick et al. (Biol Invasions, 2017) propose that the comparative functional response framework provides a unifying approach for the study of invasive species. We agree that functional responses are an important and powerful quantitative description of consumer effects on resources, and co-opting classical ecological theory to better predict invasive species impacts is a laudable move for invasion biology. However, we fear that the early successes of select examples of the comparative functional response (CFR) approach has led Dick et al. to exaggerate the generality of its utility, and about its ability to unify the field. Further, they fail to provide a convincing argument why CFR is better than existing tools such as invasion history or impact indices, even when considering emerging or potential invaders. In this response we provide details of three conceptual issues stemming from classical ecological theoretical frameworks and two practical problems that Dick et al. and other CFR proponents need to address

Sexual dimorphism in bite performance drives morphological variation in chameleons

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

Sequential Fragmentation of Pleistocene Forests in an East Africa Biodiversity Hotspot: Chameleons as a Model to Track Forest History

CITATION: Measey, G. J. & Tolley, K. A. 2011. Sequential fragmentation of Pleistocene forests in an East Africa biodiversity hotspot : chameleons as a model to track forest history. PLoS ONE, 6(10): e26606, doi:10.1371/journal.pone.0026606.The original publication is available at http://journals.plos.org/plosoneBackground The Eastern Arc Mountains (EAM) is an example of naturally fragmented tropical forests, which contain one of the highest known concentrations of endemic plants and vertebrates. Numerous paleo-climatic studies have not provided direct evidence for ancient presence of Pleistocene forests, particularly in the regions in which savannah presently occurs. Knowledge of the last period when forests connected EAM would provide a sound basis for hypothesis testing of vicariance and dispersal models of speciation. Dated phylogenies have revealed complex patterns throughout EAM, so we investigated divergence times of forest fauna on four montane isolates in close proximity to determine whether forest break-up was most likely to have been simultaneous or sequential, using population genetics of a forest restricted arboreal chameleon, Kinyongia boehmei. Methodology/Principal Findings We used mitochondrial and nuclear genetic sequence data and mutation rates from a fossil-calibrated phylogeny to estimate divergence times between montane isolates using a coalescent approach. We found that chameleons on all mountains are most likely to have diverged sequentially within the Pleistocene from 0.93–0.59 Ma (95% HPD 0.22–1.84 Ma). In addition, post-hoc tests on chameleons on the largest montane isolate suggest a population expansion ∼182 Ka. Conclusions/Significance Sequential divergence is most likely to have occurred after the last of three wet periods within the arid Plio-Pleistocene era, but was not correlated with inter-montane distance. We speculate that forest connection persisted due to riparian corridors regardless of proximity, highlighting their importance in the region's historic dispersal events. The population expansion coincides with nearby volcanic activity, which may also explain the relative paucity of the Taita's endemic fauna. Our study shows that forest chameleons are an apposite group to track forest fragmentation, with the inference that forest extended between some EAM during the Pleistocene 1.1–0.9 Ma.http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0026606Publisher's versio

Mind the gaps: investigating the cause of the current range disjunction in the Cape Platanna, Xenopus gilli (Anura: Pipidae)

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

Frog origins: inferences based on ancestral reconstructions of locomotor performance and anatomy

Frogs are the most species-rich and ecologically diverse group of amphibians and are characterized by a unique body plan including long legs, elongated ilia, and fused caudal vertebrae. Stem anurans such as Triadobatrachus or Czatkobatrachus have been suggested to have used jumping or hopping as part of their locomotor repertoire based on their anatomy. The earliest known true frog, Prosalirus bitis was suggested to have been a proficient jumper. However, data on jumping performance in frogs have never been used to attempt reconstruction of ancestral features at the base of the radiation. Here we provide data on jumping performance (forces and acceleration) in 20 species of extant frogs including representatives of most of the early radiating clades. We next use ancestral character value inferences to assess ancestral features. Our analyses suggest that frog ancestors were of small to medium size, had relatively short limbs, produced rather low jump forces, yet were capable of relatively high acceleration. Given the short limbs and low forces, the unique frog bauplan with a reduced vertebral column and a mobile ilio-sacral joint may not have been an adaptation for powerful jumping

What’s for dinner? Diet and potential trophic impact of an invasive anuran Hoplobatrachus tigerinus on the Andaman archipelago

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

Counting chirps : acoustic monitoring of cryptic frogs

Funding for the frog survey was received from the National Geographic Society/Waitt Grants Program (No. W184-11). The EPSRC and NERC helped to fund this research through a PhD grant (No. EP/1000917/1) to D.L.B. R.A. and G.J.M. acknowledge initiative funding from the National Research Foundation of South Africa.1 .  Global amphibian declines have resulted in a vital need for monitoring programmes that follow population trends. Monitoring using advertisement calls is ideal as choruses are undisturbed during data collection. However, methods currently employed by managers frequently rely on trained observers, and/or do not provide density data on which to base trends. 2 .  This study explores the utility of monitoring using acoustic spatially explicit capture-recapture (aSECR) with time of arrival (ToA) and signal strength (SS) as a quantitative monitoring technique to measure call density of a threatened but visually cryptic anuran, the Cape peninsula moss frog Arthroleptella lightfooti. 3 .  The relationships between temporal and environmental variables (date, rainfall, temperature) and A. lightfooti call density at three study sites on the Cape peninsula, South Africa were examined. Acoustic data, collected from an array of six microphones over four months during the winter breeding season, provided a time series of call density estimates. 4 .  Model selection indicated that call density was primarily associated with seasonality fitted as a quadratic function. Call density peaked mid-breeding season. At the main study site, the lowest recorded mean call density (0·160 calls m-2 min-1) occurred in May and reached its peak mid-July (1·259 calls m-2 min-1). The sites differed in call density, but also the effective sampling area. 5 .  Synthesis and applications.The monitoring technique, acoustic spatially explicit capture–recapture (aSCR), quantitatively estimates call density without disturbing the calling animals or their environment, while time of arrival (ToA) and signal strength (SS) significantly add to the accuracy of call localisation, which in turn increases precision of call density estimates without the need for specialist field staff. This technique appears ideally suited to aid the monitoring of visually cryptic, acoustically active species.Publisher PDFPeer reviewe

In a rough spot: Declines in Arthroleptella rugosa calling densities are explained by invasive pine trees

Basing conservation interventions on evidence is important for justifying their associated cost and gauging their effectiveness. For amphibians, the number of studies available to support conservation action plans is limited. Here, we sought to determine the effect of invasive pine trees on the calling densities of a Critically Endangered frog species, endemic to the Western Cape province in South Africa. The Rough Moss frog, Arthroleptella rugosa, is an anuran restricted to a small patch of fire-driven fynbos habitat prone to invasion by Cluster Pines, Pinus pinaster. We use acoustic spatially explicit capture–recapture methods to estimate frog densities at multiple sites (n = 12) over ten years. Sites were clas-sified as invaded or uninvaded by P. pinaster, and this information, along with the time since the last fire, were used as explanatory variables for frog density in a generalized linear mixed model. Frog densities were found to be significantly affected by P. pinaster invasion status

A general framework for animal density estimation from acoustic detections across a fixed microphone array

Acoustic monitoring can be an efficient, cheap, non‐invasive alternative to physical trapping of individuals. Spatially explicit capture–recapture (SECR) methods have been proposed to estimate calling animal abundance and density from data collected by a fixed array of microphones. However, these methods make some assumptions that are unlikely to hold in many situations, and the consequences of violating these are yet to be investigated. We generalize existing acoustic SECR methodology, enabling these methods to be used in a much wider variety of situations. We incorporate time‐of‐arrival (TOA) data collected by the microphone array, increasing the precision of calling animal density estimates. We use our method to estimate calling male density of the Cape Peninsula Moss Frog Arthroleptella lightfooti. Our method gives rise to an estimator of calling animal density that has negligible bias, and 95% confidence intervals with appropriate coverage. We show that using TOA information can substantially improve estimate precision. Our analysis of the A. lightfooti data provides the first statistically rigorous estimate of calling male density for an anuran population using a microphone array. This method fills a methodological gap in the monitoring of frog populations and is applicable to acoustic monitoring of other species that call or vocalize

Frog eat frog: exploring variables influencing anurophagy

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