Paul Webala
Maasai Mara University, Forestry and Wildlife Management, Faculty Member
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White or clear (“whitish”) wings are a distinct feature in about 30 species of tropical insectivorous bats (Mammalia: Chiroptera) belonging to three families (Emballonuridae, Molossidae, and Vespertilionidae). Such wings may provide... more
White or clear (“whitish”) wings are a distinct feature in about 30 species of tropical insectivorous bats (Mammalia: Chiroptera) belonging to three families (Emballonuridae, Molossidae, and Vespertilionidae). Such wings may provide camouflage against the sky at dusk and dawn, when bats commute to and from the roost and are vulnerable to aerial predation from birds. We tested this hypothesis by comparing the contrast of black, white, and transparent plastic models against the evening sky. Compared with normally dark wings, white and particularly transparent wings indeed reduce the contrast against the sky and may also prevent overheating in bats flying in daylight. Whitish wings could facilitate earlier evening emergence and later morning return, increasing access to crepuscular or diurnal insects as food. But whitish wings become maladaptive near artificial lights, where they are highly visible when illuminated against the dark sky. Pale but colored (not whitish) wings and reticulated patterns on translucent wings in some African and south Asian bats may be variations on the same theme, functional as camouflage against a lit background of vegetation and shades.
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Research Interests: Biology, Ecology, Endemism, Fauna, Rainforest, and 4 moreGenus, IUCN Red List, Threatened Species, and Taxon
The genus Myotis is nearly cosmopolitan and the second-most speciose genus of mammals, but its Afrotropical members are few and poorly known. We analyzed phylogenetic and phylogeographic relationships of six of the eight known... more
The genus Myotis is nearly cosmopolitan and the second-most speciose genus of mammals, but its Afrotropical members are few and poorly known. We analyzed phylogenetic and phylogeographic relationships of six of the eight known Afrotropical species using Cytb and sequences from four nuclear introns. Using Bayesian and maximum-likelihood approaches to generate single-locus, concatenated, and species trees, we confirmed prior evidence that the clade containing Afrotropical Myotis also contains both Palearctic and Indomalayan members. Additionally, we demonstrate that M. bocagii is sister to the Indian Ocean species M. anjouanensis, that this group is sister to M. tricolor and the Palearctic M. emarginatus, and find evidence suggesting that M. welwitschii is the earliest-diverging Afrotropical species and sister to the remainder. Although M. tricolor and M. welwitschii are both currently regarded as monotypic, both mitochondrial and nuclear data sets document significant, largely concordant geographic structure in each. Evidence for the distinction of two lineages within M. tricolor is particularly strong. On the other hand, geographic structure is lacking in M. bocagii, despite the current recognition of two subspecies in that species. Additional geographic sampling (especially at or near type localities), finer-scale sampling (especially in zones of sympatry), and integrative taxonomic assessments will be needed to better document this radiation and refine its nomenclature.
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Bats are prodigious consumers of agricultural and forest pests, and are, therefore, a natural asset for agricultural productivity, suppressing populations of such pests. This study provides baseline information of diet of 143 bats... more
Bats are prodigious consumers of agricultural and forest pests, and are, therefore, a natural asset for agricultural productivity, suppressing populations of such pests. This study provides baseline information of diet of 143 bats belonging to eight insectivorous bat species from agricultural areas of Rwanda while evaluating the effectiveness of bats as pest suppressors. Using DNA metabarcoding to analyze bat fecal pellets, 85 different insect species were detected, with 60% (n = 65), 64% (n = 11) and 78% (n = 9) found to be agricultural pests from eastern, northern and western regions, respectively. Given the high percentages of agricultural pests detected, we submit that Rwandan insectivorous bats have the capacity for biocontrol of agricultural pests. Rwandan bat populations should be protected and promoted since they may foster higher crop yields and sustainable livelihoods.
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Livestock depredation is a major conservation challenge globally, causing significant economic losses to pastoralists and threatening large carnivore species outside protected areas. Our study investigated the temporal and spatial... more
Livestock depredation is a major conservation challenge globally, causing significant economic losses to pastoralists and threatening large carnivore species outside protected areas. Our study investigated the temporal and spatial distribution of livestock depredation incidences, carnivore species associated with livestock depredation, and assessed mitigation measures in Maasai Mara Conservancies in Southern Kenya. Using daily monitoring of livestock depredation cases, we made comparisons between livestock attacks occurring in predator-proof bomas and those with traditional kraals. A total of 305 livestock depredation incidents were recorded between January and December 2021, translating to a total tally of 1411 livestock maimed or killed. Most livestock depredation incidents occurred during the day (59%) as opposed to night (41%), but this difference was not significant. Livestock depredation incidents in the nighttime occurred mostly inside traditional kraals (34%) and occurred th...
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Background and Methods: To investigate virus diversity in hot zones of probable pathogen spillover, 54 oral-fecal swabs were processed from five bat species collected from three cave systems in Kenya, using metagenome sequencing. Results:... more
Background and Methods: To investigate virus diversity in hot zones of probable pathogen spillover, 54 oral-fecal swabs were processed from five bat species collected from three cave systems in Kenya, using metagenome sequencing. Results: Viruses belonging to the Astroviridae, Circoviridae, Coronaviridae, Dicistroviridae, Herpesviridae and Retroviridae were detected, with unclassified viruses. Retroviral sequences were prevalent; 74.1% of all samples were positive, with distinct correlations between virus, site and host bat species. Detected retroviruses comprised Myotis myotis, Myotis ricketti, Myotis daubentonii and Galidia endogenous retroviruses, murine leukemia virus-related virus and Rhinolophus ferrumequinum retrovirus (RFRV). A near-complete genome of a local RFRV strain with identical genome organization and 2.8% nucleotide divergence from the prototype isolate was characterized. Bat coronavirus sequences were detected with a prevalence of 24.1%, where analyses on the ORF1a...
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Migrating grazers and carnivores respond to seasonal changes in the environment and often match peaks in resource abundance. However, it is unclear whether and how frugivorous animals use phenological events to time migration, especially... more
Migrating grazers and carnivores respond to seasonal changes in the environment and often match peaks in resource abundance. However, it is unclear whether and how frugivorous animals use phenological events to time migration, especially in the tropics. The straw‐coloured fruit bat Eidolon helvum, Africa's most gregarious fruit bat, forms large seasonal colonies throughout much of sub‐Saharan Africa. We hypothesized that aggregations of E. helvum match the timing of their migration with phenologies of plant growth or precipitation. Using monthly colony counts from across much of the species' range, we matched peak colony size to landscape phenologies and explored the variation among colonies matching the overall closest phenological event. Peak colony size was closest to the peak instantaneous rate of green‐up, and sites with closer temporal matching were associated with higher maximum greenness, short growing season and larger peak colony size. Eidolon helvum seem to time t...
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The paucity of studies on Egyptian mongoose Herpestes ichneumon (Linnaeus, 1758) in Africa highlights the need for baseline information on the geographic range of this species as well as factors that may determine its distribution. This... more
The paucity of studies on Egyptian mongoose Herpestes ichneumon (Linnaeus, 1758) in Africa highlights the need for baseline information on the geographic range of this species as well as factors that may determine its distribution. This study presents eight novel locality records of H. ichneumon in Laikipia County, central Kenya, addresses questions on the species’ distribution in Africa, and predicts environmental (climatic) suitability across its range. From a total of 4,822 H. ichneumon occurrence records, we used 4,432 georeferenced records to generate distribution maps, conduct ecological niche modelling, and identify environmental limits for this species across its range with a focus on Africa. 20% of all records originate from continental Africa, including 121 records for Kenya. Despite extensive field research and predicted habitat suitability, H. ichneumon has not been previously reported in Laikipia County. Our niche models, however, predicted parts of Laikipia to be envir...
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In the light of the urgency raised by the COVID-19 pandemic, global investment in wildlife virology is likely to increase, and new surveillance programmes will identify hundreds of novel viruses that might someday pose a threat to humans.... more
In the light of the urgency raised by the COVID-19 pandemic, global investment in wildlife virology is likely to increase, and new surveillance programmes will identify hundreds of novel viruses that might someday pose a threat to humans. To support the extensive task of laboratory characterization, scientists may increasingly rely on data-driven rubrics or machine learning models that learn from known zoonoses to identify which animal pathogens could someday pose a threat to global health. We synthesize the findings of an interdisciplinary workshop on zoonotic risk technologies to answer the following questions. What are the prerequisites, in terms of open data, equity and interdisciplinary collaboration, to the development and application of those tools? What effect could the technology have on global health? Who would control that technology, who would have access to it and who would benefit from it? Would it improve pandemic prevention? Could it create new challenges? This artic...
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Simultaneous use of domestic spaces by humans and wildlife is little understood, despite global ubiquity, and can create an interface for human exposure to wildlife pathogens. Bats are a pervasive synanthropic taxon and are associated... more
Simultaneous use of domestic spaces by humans and wildlife is little understood, despite global ubiquity, and can create an interface for human exposure to wildlife pathogens. Bats are a pervasive synanthropic taxon and are associated with several pathogens that can spill over and cause disease in humans. Urbanization has destroyed much natural bat habitat and, in response, many species increasingly use buildings as roosts. The purpose of this study was to characterize human interactions with bats in shared buildings to assess potential for human exposure to and spillover of bat-borne pathogens. We surveyed 102 people living and working in buildings used as bat roosts in Taita-Taveta county, Kenya between 2021 and 2023. We characterized and quantified the duration, intensity, and frequency of human-bat interactions occurring in this common domestic setting. Survey respondents reported living with bats in buildings year-round, with cohabitation occurring consistently for at least 10 years in 38% of cases. Human contact with bats occurred primarily through direct and indirect routes, including exposure to excrement (90% of respondents), and direct touching of bats (39% of respondents). Indirect contacts most often occurred daily, and direct contacts most often occurred yearly. Domestic animal consumption of bats was also reported (16% of respondents). We demonstrate that shared building use by bats and humans in rural Kenya leads to prolonged, frequent, and sometimes intense interactions between bats and humans, consistent with interfaces that can facilitate exposure to bat pathogens and subsequent spillover. Identifying and understanding the settings and practices that may lead to zoonotic pathogen spillover is of great global importance for developing countermeasures, and this study establishes bat roosts in buildings as such a setting.
Ecological information on wildlife reservoirs is fundamental for research targeting prevention of zoonotic infectious disease, yet basic information is lacking for many species in global hotspots of disease emergence. We provide the first... more
Ecological information on wildlife reservoirs is fundamental for research targeting prevention of zoonotic infectious disease, yet basic information is lacking for many species in global hotspots of disease emergence. We provide the first estimates of synchronicity, magnitude, and timing of seasonal birthing in Mops condylurus, a putative ebolavirus host, and a co-roosting species, Mops pumilus (formerly Chaerephon pumilus). We show that population-level synchronicity of M. condylurus birthing is wide (* 8.5 weeks) and even wider in M. pumilus (> 11 weeks). This is predicted to promote the likelihood of filovirus persistence under conditions of biannual birthing (two births per year). Ecological features underlying the magnitude of the birth pulse-relative female abundance (higher than expected for M. condylurus and lower for M. pumilus, based on literature) and reproductive rate (lower than expected)-will have countering effects on birthing magnitude. Species-specific models are needed to interpret how identified birth pulse attributes may interact with other features of molossid ebolavirus ecology to influence infection dynamics. As a common feature of wildlife species, and a key driver of infection dynamics, detailed information on seasonal birthing will be fundamental for future research on these species and will be informative for bat-borne zoonoses generally.
The file provides data on the forearm length and resting frequency echolocation calls of individuals bats used in the analyses of the paper titled "Testing the Sensory Drive Hypothesis: geographic variation in echolocation... more
The file provides data on the forearm length and resting frequency echolocation calls of individuals bats used in the analyses of the paper titled "Testing the Sensory Drive Hypothesis: geographic variation in echolocation frequencies of Geoffroy's horseshoe bat (Rhinolophidae: Rhinolophus clivosus )" published in Plos One in 2017.
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Background The recognition and delineation of morphologically indistinguishable cryptic species can have broad implications for wildlife conservation, disease ecology and accurate estimates of biodiversity. Parasites are intriguing in the... more
Background The recognition and delineation of morphologically indistinguishable cryptic species can have broad implications for wildlife conservation, disease ecology and accurate estimates of biodiversity. Parasites are intriguing in the study of cryptic speciation because unique evolutionary pressures and diversifying factors are generated by ecological characteristics of host-parasite relationships, including host specificity. Bat flies (Diptera: Nycteribiidae and Streblidae) are obligate, hematophagous ectoparasites of bats that generally exhibit high host specificity. One rare exception is Penicillidia fulvida (Diptera: Nycteribiidae), an African bat fly found in association with many phylogenetically distant hosts. One explanation for P. fulvida’s extreme polyxeny is that it may represent a complex of host-specific yet cryptic species, an increasingly common finding in molecular genetic studies of supposed generalist parasites. Methods A total of 65 P. fulvida specimens were c...
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Les peuplements de chauves-souris ont été étudiés par piégeage sur sept sites sur la bordure Est du lac Turkana, Kénya, région où le statut et la distribution de ces mammifères demeuraient inconnus. Nous avons capturé 97 individus de 13... more
Les peuplements de chauves-souris ont été étudiés par piégeage sur sept sites sur la bordure Est du lac Turkana, Kénya, région où le statut et la distribution de ces mammifères demeuraient inconnus. Nous avons capturé 97 individus de 13 espèces appartenant à 6 familles : Mégadermatidés (Cardioderma cor), Rhinolophidés (Rhinolophus clivosus), Nyctéridés (Nycteris hispida et N. thebaica), Vespertilionidés (Myotis tricolor, Nycticeinops schlieffeni, Pipistrellus kuhlii et Neoromicia nanus), Molossidés (Chaerephon pumila, Mops condylurus et M. demonstrator), Emballonuridés (Taphozous perforatus et Coleura afra). Lavia frons (Mégadermatidés) a été observé mais non capturé. La présence de bâtiments autour des stations de piégeage a affecté de manière significative le succès du piégeage. L'abondance, la richesse et la diversité se sont avérées variables selon les sites, en relation avec des différences dans la structure écologique des habitats forestiers, semi-désertiques ou insulaires...
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Understanding biodiversity patterns as well as drivers of population declines, and range losses provides crucial baselines for monitoring and conservation. However, the information needed to evaluate such trends remains unstandardised and... more
Understanding biodiversity patterns as well as drivers of population declines, and range losses provides crucial baselines for monitoring and conservation. However, the information needed to evaluate such trends remains unstandardised and sparsely available for many taxonomic groups and habitats, including the cave-dwelling bats and cave ecosystems. Here, we present the DarkCideS 1.0, a global database of bat caves and bat species based on curated data from the literature, personal collections, and existing datasets. The database contains information for geographical distribution, ecological status, species traits, and parasites and hyperparasites for 679 bat species known to occur in caves or use caves in their life-histories. The database contains 6746 georeferenced occurrences for 402 cave-dwelling bat species from 2002 cave sites in 46 countries and 12 terrestrial biomes. The database has been developed to be a collaborative, open-access, and user-friendly platform, allowing con...
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For forty years, there has been growing uncertainty about whether Hill's horseshoe bat (Rhinolophus hilli) still persists in Nyungwe National Park, Rwanda. Only known from one small area within the National Park, R. hilli is listed as... more
For forty years, there has been growing uncertainty about whether Hill's horseshoe bat (Rhinolophus hilli) still persists in Nyungwe National Park, Rwanda. Only known from one small area within the National Park, R. hilli is listed as Critically Endangered by the International Union for the Conservation of Nature (IUCN) based on its extremely small geographic range and presumed low number of mature individuals. Here, we present and describe bat species occurrence data contributed to the Global Biodiversity Information Facility (GBIF) that we collected as part of a long-term collaborative project to rediscover this lost species. This data paper describes the conservation relevance, survey methods, and findings resulting from cave roost surveys, capture surveys, and acoustic sampling of bat echolocation activity in Nyungwe National Park and surrounding areas in southwestern Rwanda from 2013-2020. We report the discovery of an extant population of Hill's horseshoe bat (Rhinolop...
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Maximum likelihood gene trees inferred for nuclear introns using IQ-TREE (Aâ D) and Bayesian gene trees inferred for nuclear introns using MRBAYES (Eâ H). Nodal support is indicated above branches. Museum acronyms are defined in... more
Maximum likelihood gene trees inferred for nuclear introns using IQ-TREE (Aâ D) and Bayesian gene trees inferred for nuclear introns using MRBAYES (Eâ H). Nodal support is indicated above branches. Museum acronyms are defined in Additional file 1. (PDF 1980 kb)
Uncorrected cyt-b p-distances among (below diagonal) and within (numbers on diagonal) African Rhinolophus clades calculated in MEGA X 10.0.5 [36]. (XLS 46 kb)
Primer information for regions amplified in the current study. (DOCX 18 kb)
List of specimens used in genetic analyses of Rhinolophus. (XLSX 55 kb)
We present data on the collections of different small mammals (mainly bats, shrews and rodents) collected from the interior of Arabuko-Sokoke Forest (ASF) and human-modified habitats (farmlands) around the forest. A total of 534 specimens... more
We present data on the collections of different small mammals (mainly bats, shrews and rodents) collected from the interior of Arabuko-Sokoke Forest (ASF) and human-modified habitats (farmlands) around the forest. A total of 534 specimens have been collected in the area from 1932-2017. Some collections of 1932-1972 were collected in areas which already had the indigenous natural vegetation typical of east African coastal biome, but currently such vegetation has completely been lost and replaced with farmlands and human settlement areas, thus not suitable for supporting the coastal species like the Golden-rumped Sengi (Rhynchocyon chrysopygus). Hence, the collection data can be used to inform on the changes in the size of ASF overtime, as result of anthropogenic activities around it and how it consequently affects the distribution of species conserved in this forest.
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In light of the urgency raised by the COVID-19 pandemic, global investment in wildlife virology is likely to increase, and new surveillance programs will identify hundreds of novel viruses that might someday pose a threat to humans. Our... more
In light of the urgency raised by the COVID-19 pandemic, global investment in wildlife virology is likely to increase, and new surveillance programs will identify hundreds of novel viruses that might someday pose a threat to humans. Our capacity to identify which viruses are capable of zoonotic emergence depends on the existence of a technology—a machine learning model or other informatic system—that leverages available data on known zoonoses to identify which animal pathogens could someday pose a threat to global health. We synthesize the findings of an interdisciplinary workshop on zoonotic risk technologies to answer the following questions: What are the prerequisites, in terms of open data, equity, and interdisciplinary collaboration, to the development and application of those tools? What effect could the technology have on global health? Who would control that technology, who would have access to it, and who would benefit from it? Would it improve pandemic prevention? Could it...
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Despite being nearly 10 months into the COVID-19 (coronavirus disease 2019) pandemic, the definitive animal host for SARS-CoV-2 (severe acute respiratory syndrome coronavirus 2), the causal agent of COVID-19, remains unknown.... more
Despite being nearly 10 months into the COVID-19 (coronavirus disease 2019) pandemic, the definitive animal host for SARS-CoV-2 (severe acute respiratory syndrome coronavirus 2), the causal agent of COVID-19, remains unknown. Unfortunately, similar problems exist for other betacoronaviruses, and no vouchered specimens exist to corroborate host species identification for most of these pathogens. This most basic information is critical to the full understanding and mitigation of emerging zoonotic diseases. To overcome this hurdle, we recommend that host-pathogen researchers adopt vouchering practices and collaborate with natural history collections to permanently archive microbiological samples and host specimens. Vouchered specimens and associated samples provide both repeatability and extension to host-pathogen studies, and using them mobilizes a large workforce (i.e., biodiversity scientists) to assist in pandemic preparedness. We review several well-known examples that successfull...
Research Interests: Microbiology, Museum Education, Disease ecology, Virology, Tropical Medicine, and 14 moreNatural History, Public Health, Biodiversity, Infectious Diseases, Emerging Infectious Diseases, Museums, Natural history collections, One Health, Voucher Specimens, Natural History Museums, Host Pathogen Interactions, Mbio, Coronavirus COVID-19, and COVID-19 PANDEMIC
Vespertilionidae (class Mammalia) constitutes the largest family of bats, with ~500 described species. Nonetheless, the systematic relationships within this family are poorly known, especially among the pipistrelle-like bats of the tribes... more
Vespertilionidae (class Mammalia) constitutes the largest family of bats, with ~500 described species. Nonetheless, the systematic relationships within this family are poorly known, especially among the pipistrelle-like bats of the tribes Vespertilionini and Pipistrellini. Perhaps as a result of their drab pelage and lack of obvious morphological characters, the genus and species limits of pipistrelle-like bats remain poorly resolved, particularly in Africa, where more than one-fifth of all vesper bat species occur. Further exacerbating the problem is the accelerating description of new species within these groups. In this study, we attempt to resolve the systematic relationships among the pipistrelle-like bats of sub-Saharan Africa and Madagascar and provide a more stable framework for future systematic efforts. Our systematic inferences are based on extensive genetic and morphological sampling of > 400 individuals covering all named genera and the majority of described African ...
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Rhinonycteridae (trident bats) are a small Palaeotropical family of insectivorous bats allied to Hipposideridae. Their taxonomy has been in a state of flux. Here, we use mitochondrial and nuclear sequences to evaluate species... more
Rhinonycteridae (trident bats) are a small Palaeotropical family of insectivorous bats allied to Hipposideridae. Their taxonomy has been in a state of flux. Here, we use mitochondrial and nuclear sequences to evaluate species relationships, confirming the monophyly of both Triaenops and Paratriaenops. Although most Triaenops afer specimens are recovered as a group, mitochondrial analyses strongly support some Kenyan individuals as members of Triaenops persicus. Analyses of four nuclear introns (ACOX2, COPS7A, RODGI and STAT5A) strongly support the mitochondrial topology. Morphometric analysis of the skull, external morphology and echolocation calls confirm that the Triaenops from the Rift Valley in Kenya (Nakuru, Baringo and Pokot counties) are distinct from typical T. afer in coastal (Kilifi and Kwale counties) or interior (Laikipia and Makueni counties) colonies. We interpret these analyses to indicate that two species of Triaenops occur in East Africa: T. afer in coastal regions ...
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The Old World leaf-nosed bats (Hipposideridae) are aerial and gleaning insectivores that occur throughout the Paleotropics. Both their taxonomic and phylogenetic histories are confused. Until recently, the family included genera now... more
The Old World leaf-nosed bats (Hipposideridae) are aerial and gleaning insectivores that occur throughout the Paleotropics. Both their taxonomic and phylogenetic histories are confused. Until recently, the family included genera now allocated to the Rhinonycteridae and was recognized as a subfamily of Rhinolophidae. Evidence that Hipposideridae diverged from both Rhinolophidae and Rhinonycteridae in the Eocene confirmed their family rank, but their intrafamilial relationships remain poorly resolved. We examined genetic variation in the Afrotropical hipposiderids Doryrhina, Hipposideros, and Macronycteris using relatively dense taxon-sampling throughout East Africa and neighboring regions. Variation in both mitochondrial (cyt-b) and four nuclear intron sequences (ACOX2, COPS, ROGDI, STAT5) were analyzed using both maximum likelihood and Bayesian inference methods. We used intron sequences and the lineage delimitation method BPP—a multilocus, multi-species coalescent approach—on suppo...
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Supplementary material 3 from: Patterson BD, Webala PW, Lavery TH, Agwanda BR, Goodman SM, Kerbis Peterhans JC, Demos TC (2020) Evolutionary relationships and population genetics of the Afrotropical leaf-nosed bats (Chiroptera, Hipposideridae). ZooKeys 929: 117-161. https://doi.org/10.3897/zookey...more
Supplementary material 1 from: Patterson BD, Webala PW, Lavery TH, Agwanda BR, Goodman SM, Kerbis Peterhans JC, Demos TC (2020) Evolutionary relationships and population genetics of the Afrotropical leaf-nosed bats (Chiroptera, Hipposideridae). ZooKeys 929: 117-161. https://doi.org/10.3897/zookey...more
Supplementary material 2 from: Patterson BD, Webala PW, Lavery TH, Agwanda BR, Goodman SM, Kerbis Peterhans JC, Demos TC (2020) Evolutionary relationships and population genetics of the Afrotropical leaf-nosed bats (Chiroptera, Hipposideridae). ZooKeys 929: 117-161. https://doi.org/10.3897/zookey...more