is growing evidence that cryptic lineages exist in many morphologically conservative species of bats. It is, however, challenging to assign a meaningful taxonomic rank to these lineages, as species delineation is often complicated by conflicting data sets. Conflicting data sets may result from different types of characters (e.g. phenotypic versus genotypic) or even from different compartments of the genotype (e.g. mitochondrial versus nuclear markers). Historical, demographic or selective factors may be invoked to explain discrepancies, but final decisions regarding the species status of each lineage are often dependent on appropriate geographic and genetic sampling.The cryptic myotis (Myotis crypticus) is a typical example where morphological characters can barely differentiate it from its sister species the Natterer's bat (M. nattereri) yet mitochondrial and nuclear markers apparently show only very localized evidence of ongoing introgression between them. We used an array of 11 highly variable microsatellite markers genotyped in 320 individuals from 22 populations sampled across Western Europe to investigate more precisely population structure and the porosity of interspecific barriers in this species complex. Results based on Bayesian and multivariate assignment of individual nuclear genotypes suggest that M. crypticus and M. nattereri are hardly connected by recent gene flow, confirming their status as evolutionarily distinct species. However, local mitonuclear discordance indicates that ancient introgression occurred when these formerly allopatric species entered secondary contact during postglacial range expansions across the Alps. At the intraspecific level, Natterer's bat (M. nattereri) exhibits low genetic structuring across Western Europe, consistent with prior evidence of recent population expansion. In contrast, populations of Cryptic Myotis display pronounced western and eastern genetic differentiation reflecting the influence of Iberian and Apennine glacial refugia from which these bats likely recolonized more northerly regions.
When one or several classes are much less prevalent than another class (unbalanced data), class error rates and variable importances of the machine learning algorithm random forest can be biased, particularly when sample sizes are smaller, imbalance levels higher, and effect sizes of important variables smaller. Using simulated data varying in size, imbalance level, number of true variables, their effect sizes, and the strength of multicollinearity between covariates, we evaluated how eight versions of random forest ranked and selected true variables out of a large number of covariates despite class imbalance. The version that calculated variable importance based on the area under the curve (AUC) was least adversely affected by class imbalance. For the same number of true variables, effect sizes, and multicollinearity between covariates, the AUC variable importance ranked true variables still highly at the lower sample sizes and higher imbalance levels at which the other seven versions no longer achieved high ranks for true variables. Conversely, using the Hellinger distance to split trees or downsampling the majority class already ranked true variables lower and more variably at the larger sample sizes and lower imbalance levels at which the other algorithms still ranked true variables highly. In variable selection, a higher proportion of true variables were identified when covariates were ranked by AUC importances and the proportion increased further when the AUC was used as the criterion in forward variable selection. In three case studies, known species–habitat relationships and their spatial scales were identified despite unbalanced data.
Context Habitat suitability models (HSM) can improve our understanding of a species’ ecology and are valuable tools for informing landscape-scale decisions. We can increase HSM predictive accuracy and derive more realistic conclusions by taking a multi-scale approach. However, this process is often statistically complex and computationally intensive. Objectives We provide an easily implemented, flexible framework for sequential multi-level, multi-scale HSM and compare it to two other commonly-applied approaches: single-level, multi-scale HSM and their post-hoc combinations. Methods Our framework implements scale optimisation and model tuning at each level in turn, from the highest (population range) to the lowest (e.g. foraging habitat) level, whilst incorporating output habitat suitability indices from a higher level as a predictor. We used MaxEnt and a species of conservation concern in Britain, the lesser horseshoe bat ( Rhinolophus hipposideros ), to demonstrate and compare multi-scale approaches. Results Integrating models across levels, either by applying our framework, or by multiplying single-level model predictions, improved predictive performance over single-level models. Moreover, differences in the importance and direction of the species-environment associations highlight the potential for false inferences from single-level models or their post-hoc combinations. The single-level summer range model incorrectly identified a positive influence of heathland cover, whereas sequential multi-level models made biological sense and underlined this species’ requirement for extensive broadleaf woodland cover, hedgerows and access to buildings for roosting in rural areas. Conclusions We conclude that multi-level HSM appear superior to single-level, multi-scale approaches; models should be sequentially integrated across levels if information on species-environment relationships is of importance.
Generating, collating and using scientific evidence is key to effective conservation. We illustrate this with examples from medicine and conservation, and in more depth with an example from our own studies, in which we assess the methods (overpasses and underpasses) used to reduce habitat fragmentation and mortality of bats caused by roads. Evidence is defined as the results of quantitative tests of interventions that directly address conservation effectiveness against clearly stated goals. The results show that methods used in the past were rarely tested quantitatively and when tested were rarely effective. We discuss why it can be difficult to get scientific evidence accepted and used routinely in conservation policy and practice. Finally, we discuss how evidence gathering and use can be improved in mitigation projects.
We used capture (mist-netting) and acoustic methods to compare the species richness, abundance, and composition of a bat assemblage in different habitats in the Western Ghats of India. In the tropics, catching bats has been more commonly used as a survey method than acoustic recordings. In our study, acoustic methods based on recording echolocation calls detected greater bat activity and more species than mist-netting. However, some species were detected more frequently or exclusively by capture. Ideally, the two methods should be used together to compensate for the biases in each. Using combined capture and acoustic data, we found that protected forests, forest fragments, and shade coffee plantations hosted similar and diverse species assemblages, although some species were recorded more frequently in protected forests. Tea plantations contained very few species from the overall bat assemblage. In riparian habitats, a strip of forested habitat on the river bank improved the habitat for bats compared to rivers with tea planted up to each bank. Our results show that shade coffee plantations are better bat habitat than tea plantations in biodiversity hotspots. However, if tea is to be the dominant land use, forest fragments and riparian corridors can improve the landscape considerably for bats. We encourage coffee growers to retain traditional plantations with mature native trees, rather than reverting to sun grown coffee or coffee shaded by a few species of timber trees.
The responses of bats to land-use change have been extensively studied in temperate zones and the neotropics, but little is known from the palaeotropics. Effective conservation in heavily-populated palaeotropical hotspots requires a better understanding of which bats can and cannot survive in human-modified landscapes. We used catching and acoustic transects to examine bat assemblages in the Western Ghats of India, and identify the species most sensitive to agricultural change. We quantified functional diversity and trait filtering of assemblages in forest fragments, tea and coffee plantations, and along rivers in tea plantations with and without forested corridors, compared to protected forests.Functional diversity in forest fragments and shade-grown coffee was similar to that in protected forests, but was far lower in tea plantations. Trait filtering was also strongest in tea plantations. Forested river corridors in tea plantations mitigated much of the loss of functional diversity and the trait filtering seen on rivers in tea plantations without forested corridors. The bats most vulnerable to intensive agriculture were frugivorous, large, had short broad wings, or made constant frequency echolocation calls. The last three features are characteristic of forest animal-eating species that typically take large prey, often by gleaning.Ongoing conservation work to restore forest fragments and retain native trees in coffee plantations should be highly beneficial for bats in this landscape. The maintenance of a mosaic landscape with sufficient patches of forest, shade-grown coffee and riparian corridors will help to maintain landscape wide functional diversity in an area dominated by tea plantations.
Salim Ali’s Fruit Bat Latidens salimalii is an IUCN Red listed Endangered species known only from a few locations in southern India. Here we report three records of Latidens salimalii from the Valparai plateau and Anamalai Tiger Reserve in Tamil Nadu where this species has not been previously recorded. This bat was caught in riparian habitats close to or inside intact tropical wet forest in the Western Ghats.
Diurnal and seasonal increases in body mass and seasonal reductions in wing area may compromise a bird's ability to escape, as less of the power available from the flight muscles can be used to accelerate and elevate the animal's centre of mass. Here we investigated the effects of intra-specific variation in wing morphology on escape take-off performance in blue tits (Cyanistes caeruleus). Flights were recorded using synchronised high-speed video cameras and take-off performance was quantified as the sum of the rates of change of the kinetic and potential energies of the centre of mass. Individuals with a lower wing loading, WL (WL=body weight/ wing area) had higher escape take-off performance, consistent with the increase in lift production expected from relatively larger wings. Unexpectedly, it was found that the total power available from the flight muscles (estimated using an aerodynamic analysis) was inversely related to WL. This could simply be because birds with a higher WL have relatively smaller flight muscles. Alternatively or additionally, variation in the aerodynamic load on the wing resulting from differences in wing morphology will affect the mechanical performance of the flight muscles via effects on the muscle's length trajectory. Consistent with this hypothesis is the observation that wing beat frequency and relative downstroke duration increase with decreasing WL; both are factors that are expected to increase muscle power output. Understanding how wing morphology influences take-off performance gives insight into the potential risks associated with feather loss and seasonal and diurnal fluctuations in body mass.
To conserve biodiversity it is imperative that we understand how different species respond to land use change, and determine the scales at which habitat changes affect species' persistence. We used habitat suitability models (HSMs) at spatial scales from 100–4000m to address these concerns for bats in the Western Ghats of India, a biodiversity hotspot of global importance where the habitat requirements of bats are poorly understood. We used acoustic and capture data to build fine scale HSMs for ten species (Hesperoptenus tickelli, Miniopterus fuliginosus, Miniopterus pusillus, Myotis horsfieldii, Pipistrellus ceylonicus, Megaderma spasma, Hipposideros pomona, Rhinolophus beddomei, Rhinolophus indorouxii and Rhinolophus lepidus) in a tea-dominated landscape. Small (100–500m) scale habitat variables (e.g. percentage tea plantation cover) and distances to habitat features (e.g. distance to water) were the strongest predictors of bat occurrence, likely due to their high mobility, which enables them to exploit even small or isolated foraging areas. Most species showed a positive response to coffee plantations grown under native shade and to forest fragments, but a negative response to more heavily modified tea plantations. Two species were never recorded in tea plantations. This is the first study of bats in tea plantations globally, and the first ecological Old World bat study to combine acoustic and capture data. Our results suggest that although bats respond negatively to tea plantations, tea-dominated landscapes that also contain forest fragments and shade coffee can nevertheless support many bat species.
Bats are long-lived mammals with low reproductive rates, making them susceptible to developments that reduce reproductive output or increase mortality. Roads destroy. degrade and fragment habitats, reducing the ability of bats to roost, feed and reproduce. Current mitigation techniques have not been proven to be effective at conserving bats at the population level.34.1 Road effects vary among species depending on flight style and habitat use, but many species are affected.34.2 Roads can act as barriers to the movement of bats, and many species suffer traffic mortality attempting to cross roads.34.3 Artificial light deters some bat species while attracting others, and a policy of no lighting' is recommended for bats.34.4 Traffic noise may reduce the flight activity and foraging efficiency of bats.34.5 Protection of roosts and foraging sites during and after roadworks is critical to bat survival.34.6 Underpasses can effectively reduce the barrier effect and reduce the number of roadkills for some bat species.34.7 Other attempts to reduce the barrier and mortality effects for bats are unproven, and further research is required before widespread implementation.It is important to conduct thorough pre-construction bat surveys and bring objectivity and rigour into the design and testing of mitigation features.
Conservation increasingly operates at the landscape scale. For this to be effective, we need landscape scale information on species distributions and the environmental factors that underpin them. Species records are becoming increasingly available via data centres and online portals, but they are often patchy and biased. We demonstrate how such data can yield useful habitat suitability models, using bat roost records as an example. We analysed the effects of environmental variables at eight spatial scales (500 m - 6 km) on roost selection by eight bat species (Pipistrellus pipistrellus, P. pygmaeus, Nyctalus noctula, Myotis mystacinus, M. brandtii, M. nattereri, M. daubentonii, and Plecotus auritus) using the presence-only modelling software MaxEnt. Modelling was carried out on a selection of 418 data centre roost records from the Lake District National Park, UK. Target group pseudoabsences were selected to reduce the impact of sampling bias. Multi-scale models, combining variables measured at their best performing spatial scales, were used to predict roosting habitat suitability, yielding models with useful predictive abilities. Small areas of deciduous woodland consistently increased roosting habitat suitability, but other habitat associations varied between species and scales. Pipistrellus were positively related to built environments at small scales, and depended on large-scale woodland availability. The other, more specialist, species were highly sensitive to human-altered landscapes, avoiding even small rural towns. The strength of many relationships at large scales suggests that bats are sensitive to habitat modifications far from the roost itself. The fine resolution, large extent maps will aid targeted decision-making by conservationists and planners. We have made available an ArcGIS toolbox that automates the production of multi-scale variables, to facilitate the application of our methods to other taxa and locations. Habitat suitability modelling has the potential to become a standard tool for supporting landscape-scale decision-making as relevant data and open source, user-friendly, and peer-reviewed software become widely available.
From their earliest origins, fishes have developed a suite of adaptations for locomotion in water, which determine performance and ultimately fitness. Even without data from behaviour, soft tissue and extant relatives, it is possible to infer a wealth of palaeobiological and palaeoecological information. As in extant species, aspects of gross morphology such as streamlining, fin position and tail type are optimized even in the earliest fishes, indicating similar life strategies have been present throughout their evolutionary history. As hydrodynamical studies become more sophisticated, increasingly complex fluid movement can be modelled, including vortex formation and boundary layer control. Drag-reducing riblets ornamenting the scales of fast-moving sharks have been subjected to particularly intense research, but this has not been extended to extinct forms. Riblets are a convergent adaptation seen in many Palaeozoic fishes, and probably served a similar hydrodynamic purpose. Conversely, structures which appear to increase skin friction may act as turbulisors, reducing overall drag while serving a protective function. Here, we examine the diverse adaptions that contribute to drag reduction in modern fishes and review the few attempts to elucidate the hydrodynamics of extinct forms.