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Herpetologica, 76(2), 2020, 201–215 Ó 2020 by The Herpetologists’ League, Inc.

Chytrid Pathogen (Batrachochytrium dendrobatidis) in African : A Continental Analysis of Occurrences and Modeling of Its Potential Distribution

1,19 2 3,4 5,6 7 8 BREDA M. ZIMKUS ,VOJTECH BALA´ˇZ ,ANAT M. BELASEN ,RAYNA C. BELL ,ALAN CHANNING ,JOSEPH DOUMBIA ,ERIC B. 9 10 11 12,13 14 15 FOKAM ,LEGRAND N. GONWOUO ,ELI GREENBAUM ,VA´ CLAV GVOZDˇ IK´ ,MAREIKE HIRSCHFELD ,KATE JACKSON ,TIMOTHY 4 16 4 17 14 Y. JAMES ,CHIFUNDERA KUSAMBA ,JOANNA G. LARSON ,LISE-BETHY MAVOUNGOU ,MARK-OLIVER RO¨ DEL ,ANGE-GHISLAIN 17 14,18 ZASSI-BOULOU , AND JOHANNES PENNER 1 Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA 2 University of Veterinary and Pharmaceutical Sciences Brno, Faculty of Veterinary Hygiene and Ecology, Department of Biology and Wildlife Diseases, 612 42 Brno, Czech Republic 3 Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, NY 14853, USA 4 Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI 48109, USA 5 Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA 6 Herpetology Department, California Academy of Sciences, San Francisco, CA 94118, USA 7 Unit for Environmental Sciences and Management, North-West University, Potchefstroom, 2520, South Africa 8 ONG EnviSud Guinee,´ Commune Ratoma 030BP:558 4720 Conakry, Guinee´ 9 Department of Zoology and Physiology, University of Buea, 00237, 10 Laboratory of Zoology, Faculty of Sciences, University of Yaounde´ I, P.O. Box 812, Yaounde,´ Cameroon 11 Department of Biological Sciences, University of Texas at El Paso, 500 W. University Ave., El Paso, TX 79968, USA 12 Institute of Vertebrate Biology of the Czech Academy of Sciences, 603 65 Brno, Czech Republic 13 Department of Zoology, National Museum, 193 00 Prague, Czech Republic 14 Museum fur ¨ Naturkunde Berlin, Leibniz Institute for Evolution and Science, Invalidenstr. 43, 10115 Berlin, Germany 15 Department of Biology, Whitman College, 345 Boyer Avenue, Walla Walla, WA 99362, USA 16 Centre de Recherche en Sciences Naturelles, Departement´ de Biologie, Lwiro, The Democratic 17 Institut National de Recherche en Sciences Exactes et Naturelles (IRSEN), Cite´ Scientifique (Ex-ORSTOM), 2400, Republique´ du Congo 18 Chair of Wildlife Ecology and Wildlife Management, University of Freiburg, Tennenbacher Straße 4, 79106 Freiburg, Germany

ABSTRACT: The fungal pathogen Batrachochytrium dendrobatidis (Bd) is implicated in global declines of populations and has been documented in African specimens originally collected as far back as the 1930s. Numerous recent surveys focusing on regional pathogen prevalence have greatly increased the number of known occurrences of Bd in African species, but few studies have focused on continental distribution patterns. We analyzed all known positive occurrences of Bd in African amphibians to date, including newly reported data from Cameroon, the Democratic Republic of the Congo, , Namibia, and the Republic of the Congo. Records from both Namibia and the Republic of the Congo reported herein represent first positive occurrences for these countries. With this most comprehensive sampling of the African continent to date we identified patterns of Bd-positive occurrences associated with (1) location (i.e., biogeographic region, country), (2) , (3) life history, and (4) threat of extinction. We used fine-grained (30 arc seconds) environmental niche models (ENMs) to predict the continental distribution of Bd and identify hotspots for the pathogen, including areas not previously modeled to have high suitability for the fungus, and areas of high amphibian biodiversity from which Bd has not yet been documented. Our ENMs predicted that the environmentally suitable range of Bd encompasses vast areas of high amphibian biodiversity, including the Congo Basin and the Albertine Rift. Although our ENMs indicated that West Africa is environmentally suitable for Bd, the fungus has not been reported west of the Dahomey Gap. Likewise, the ENMs also identified regions across the Congo Basin and coastal that are environmentally suitable for the pathogen but from which Bd has not yet been reported, underscoring a need for Bd surveys in these regions. Although amphibian declines in Africa have not been directly attributed to chytridiomycosis, Bd has been detected in over one fifth of the most-threatened African amphibians. Given the presence of the hypervirulent Bd global panzootic lineage (BdGPL) in Africa, we believe that the threat of Bd as a novel pathogen may be underestimated and that focused research is urgently needed to identify which species are susceptible to Bd-driven declines. Key words: Chytridiomycosis; Fungus; Habitat; Life history; Maxent; Taxonomy

AMPHIBIANS are currently the most threatened vertebrate der Sluijs et al. 2016; Stegen et al. 2017) and causing steep group, with approximately one third of all species considered declines in populations of Fire Salamanders (Salamandra in danger of extinction with numbers continuing to rise salamandra) in the Netherlands, Belgium, and Germany. (Stuart et al. 2004; Wake and Vrendenberg 2008; IUCN Recently, Bsal was revealed to induce mass mortality in 2019). A number of causes have been linked to global Marbled Newts (Triturus marmoratus) in north-eastern declines of amphibian populations including chytridiomyco- Spain (Martel et al. 2020). Batrachochytrium dendrobatidis sis, an infectious disease caused by the chytrid fungus has been detected in 41% of amphibian species across 63% Batrachochytrium dendrobatidis (Bd), which infects the of the countries in which sampling has been reported, and keratin-containing epidermis in adults and the mouthparts in chytridiomycosis has contributed to the decline of an tadpoles (Berger et al. 1998, 2005; Johnson 2006). A second estimated 501 amphibian species worldwide, representing a species of chytrid, B. salamandrivorans (Bsal), has more catastrophic loss to biodiversity and the greatest loss recently been identified and affects salamanders, inflicting attributable to a pathogen to date (Scheele et al. 2019). skin ulcerations and necrosis (Martel et al. 2013; Spitzen-van The greatest losses attributed to Bd have been found in Central and South America, especially in large-bodied, 19 CORRESPONDENCE: email, [email protected] range-restricted anurans in wet climates, whereas the status

201 202 Herpetologica 76(2), 2020 of African amphibian chytridiomycosis-associated declines is 2019). However, the recent identification of the hyperviru- largely unknown (Hirschfeld et al. 2016; Scheele et al. 2019). lent Bd global panzootic lineage (BdGPL) in African Soto-Azat et al. (2010) documented the oldest known amphibians suggests that the threat of Bd as a novel African case of chytrid infection at the time, having detected pathogen may be underestimated (O’Hanlon et al. 2018; Bd from a museum specimen of Xenopus fraseri collected Byrne et al. 2019; Doherty-Bone et al. 2019). Herein, we from Cameroon in 1933. They suggested that southern analyze all known positive occurrences in African amphib- Africa may be the source of Bd, attributing the global spread ians to date, which include previously unpublished occur- of the fungus to the use of Xenopus laevis in human rences of Bd in several countries, to identify patterns pregnancy assays, beginning in the 1930s, and subsequent associated with distribution, taxonomic group, life history, development as a model organism for biomedical research and conservation status. We employ environmental niche (Gurdon and Hopwood 2000; Weldon et al. 2004, 2007). modeling (ENM) to test the hypothesis that areas across the Xenopus laevis is a subclinical carrier of Bd infection, and Central Africa subregion that were not previously modeled data suggest this species has exhibited a steady prevalence of as suitable habitat for Bd, specifically within the Congo infections for over 8 decades (Weldon et al. 2004; Skerratt et Basin, will be predicted as environmentally suitable for the al. 2007; Vrendenberg et al. 2013). O’Hanlon et al. (2018) fungus, given new occurrence data in this region. later refuted the ‘‘Out of Africa’’ hypothesis, concluding that the pathogen likely emerged from East Asia at the beginning MATERIALS AND METHODS of the 20th Century. The study identified known lineages of Positive Occurrence Analyses the fungus via the analysis of the genomes of hundreds of Bd samples collected from wild and captive amphibians in A systematic review of all known positive occurrences of Australia, North and South America, Africa, Asia, and Bd in continental Africa was conducted, including both Europe. They concluded that samples from Korea are published records and new accounts. Methods used to report descended from the ancestral population that gave rise to the new positive occurrences from Namibia (2007), Cameroon pandemic strain. (2009, 2016), the Democratic Republic of the Congo (2009, Although Africa was once the hypothesized source of Bd, 2010, 2013), Gabon (2013), and the Republic of the Congo the taxonomic and geographic prevalence of the pathogen (2008, 2010) are included in the Supplemental Materials and across the continent is still poorly documented. Predictions Methods (available online). Data from these new occurrenc- regarding the distribution of Bd in Africa were first based on es and previously published accounts were compiled to models made using Bd-positive localities in the New World include country, site coordinates (latitude, longitude), (Ron 2005). Although the known African localities fell within elevation, host (order, family, , species), date (year), highly suitable regions, few positive occurrences, including number positive (number of specimens of the species from only 11 in South Africa and a single site in Kenya, were the site testing positive), method(s) used to detect Bd, strain available to robustly test the model. Multiple studies have if identified with genome sequencing or genotyping assay, since assessed the global risk of Bd infection to amphibians and original citation if data were previously published. using environmental niche modeling, finding that the Original authors were contacted to obtain coordinate data if geographic ranges of declining species overlapped with the this information was not reported in the corresponding highest environmental suitability for the fungus, and further publication, and records were updated accordingly. Local- linking rapid amphibian declines with the emergence of Bd ities were assigned to one of the seven major biogeographic (Lotters¨ et al. 2009; Rodder¨ et al. 2009). The highest areas of areas in Africa that were previously used by Zimkus et al. suitability for Bd were in temperate and subtropical regions (2017): (1) Northern Africa (includes both Sahel and as well as montane regions in the tropics. Risk assessments Saharan zones; Linder et al. 2012), (2) West Africa (west concluded that only 8% of those species with the highest of the Cross River in eastern near the border with threat of extinction were African endemics, and none of Cameroon; Penner et al. 2011), (3) Central Africa (core of these had tested positive with Bd in nature (Rodder¨ et al. the continent), including the Gulf of Guinea Islands (Measey 2009). Penner et al. (2013) produced the first fine-scale et al. 2007; Bell et al. 2015), (4) Albertine Rift (including continental environmental niche model for Africa, identify- eastern Democratic Republic of the Congo and western ing fewer areas of suitability for Bd, excluding regions of Burundi, Rwanda and Uganda; Linder et al. 2012), (5) East Angola, Namibia, and Zambia that had been predicted by the Africa (Kenya, eastern Uganda, eastern Burundi, eastern two previous studies. Widespread sampling in West Africa Rwanda, Tanzania, Malawi, northern Mozambique, Zambia), also suggested that the fungus had not traveled west of the (6) Horn of Africa (includes Ethiopian and Somalian zones; Dahomey Gap, a natural savanna corridor separating the Linder et al. 2012), and (7) Southern Africa (south of the Upper and Lower Guinean rainforest blocks (Penner et al. Zambezi and Cunene Rivers; Leache´ et al. 2014). Scientific 2013). names were updated according to Frost (2020). Publication Regional herpetological surveys conducted in the past date was used if collection date was not reported. If positive decade have greatly increased the number of known Bd occurrences were reported for a multi-year study and the occurrences in African amphibians, and a number of studies specific year could not be determined, the first year reported have also tested museum specimens to elucidate the historic was used in analyses. Method(s) used to detect the presence prevalence of the pathogen in various African regions of Bd were reported using four different categories: (1) (Seimon et al. 2015; Hirschfeld et al. 2016; Hydeman et al. Morphology or histology—includes morphological examina- 2017). Relative to other parts of the world, our understand- tion of tadpole mouthparts (e.g., visualization of whole- ing of Bd prevalence and chytridiomycosis-associated mount preparations under microscope, use of photographs amphibian declines is very limited in Africa (Scheele et al. of the mouth taken in the field) or histological examination of ZIMKUS ET AL.—CHYTRID IN AFRICAN AMPHIBIANS 203 paraffin-embedded and sectioned skin tissue or toe clips olds from all ENMs was used for the average ENMs. (possibly from museum specimens) that were stained to Validation of ENMs was conducted using area under the identify Bd zoosporangia with zoospores; (2) polymerase curve (AUC) values and refers to the receiver operating chain reaction (PCR) or quantitative real-time Taqman PCR characteristic (ROC) curve, which is threshold-independent (qPCR) from museum specimens—PCR or qPCR diagnostic and commonly used (e.g., Elith et al. 2006). assay using Bd primers of preserved museum specimens fixed in various ways (e.g., formalin-fixed, industrial meth- RESULTS ylated spirits, preservation method unknown); (3) PCR/ Analysis of Positive Occurrence Data qPCR from field samples—PCR or qPCR diagnostic assay using Bd primers of skin swabs or toe clips taken in field and/ The total number of African localities with confirmed Bd or excision of mouthparts from newly collected tadpoles; and presence by species (n ¼ 1084; Supplemental Material S1) (4) Genome sequencing or genotyping—assays used to represents positive occurrences in 2753 specimens, including identify the particular strain of Bd. If multiple methods 18 families and 47 genera of , as well as 4 families and 7 were used, the method providing the most genomic genera of (Supplemental Material S1). Additional information was used. We tested for significant relationships information about 259 new positive occurrences are reported between Bd-positive occurrences and lifetime average herein, including the first records from Namibia and the aquatic index for amphibian families and genera using linear Republic of the Congo (see Supplemental Material S3 in the regression. Adult and larval habitat were coded for all Supplemental Materials available online). Batrachochytrium described species identified with Bd using Channing and dendrobatidis has been detected consistently in species of Rodel¨ (2019), and an average lifetime aquatic index ranging African amphibians from every decade since 1933 (from from 1 to 3 was calculated according to the criteria outlined museum specimens) with increased detection (and attention) by Lips et al. (2003): (1) exclusively terrestrial, (2) occupying in the last 2 decades (Fig. 1). The majority of records (76%) ponds or multiple habitats, or (3) exclusively aquatic. We were derived from PCR or qPCR analyses of field samples determined the threat level for all species identified with Bd, while far fewer were from identification using gross identifying those as extinct in the wild, critically endangered, morphology or histology (14%), testing of museum speci- or endangered on the International Union for Conservation mens (6%), or genomic sequencing or genotyping (3%). The of Nature (IUCN) Red List as of the highest concern (IUCN specific strain of Bd has been determined for 86 specimens 2019). Graphs were produced using the ggplot2 function in with the Bd Cape lineage (BdCAPE) identified in Cameroon RStudio v1.2.1335 (Wickham 2016; R Core Team 2019). and South Africa, the hypervirulent BdGPL strain identified from seven countries (Burundi, Ethiopia, Kenya, South Environmental Niche Modeling Africa, Uganda, , and Sa˜o Tome´ and Environmental niche modeling (ENM; or species distri- Prıncipe),´ and hybrid lineages identified from Burundi and bution modeling) uses occurrence data in conjunction with South Africa (Fig. 1; Supplemental Material S1). Batracho- environmental data to produce a correlative model of the chytrium dendrobatidis has been reported from amphibians environmental conditions that meet a species’ ecological originating from all six biogeographic zones, with the most requirements and to predict the relative suitability of habitat records reported from Central Africa and southern Africa in space (Peterson et al. 2011). The total number of localities (Fig. 2A inset). This is not surprising given that the greatest with confirmed Bd presences from the African continent (n number of publications have reported Bd-positive occur- ¼ 1084; see Supplemental Material S1 in the Supplemental rences from countries in these two regions (Central Africa, Materials available online) were reduced by excluding 17; southern Africa, 17; Supplemental Material S1). We occurrences without coordinate data (n ¼ 11) and either report positive occurrences from 21 countries, with the identical occurrences or those that fell within the same 1-km greatest number of records originating from South Africa, grid cell (n ¼ 452). Thus, a total of 490 unique records were Cameroon, Nigeria, and Kenya (Fig. 2A). used in ENM analyses. We used the software Maxent v3.4.1 Among the 17 families of frogs, the , (Phillips et al. 2019); a detailed description of its algorithm is , and included the highest provided by Elith et al. (2011). We acknowledge that every number of positive occurrence records (Fig. 2B). By algorithm has limitations (Qiao et al. 2015), yet Maxent has contrast, Bd is presently reported in only one or two become the standard for correlative ENMs based on the individuals in several families including Alytidae, Brevicipi- ease of its use, strong performance in comparative studies, tidae, Dicroglossidae, Microhylidae, Hylidae, and Pelobati- and success under a large variety of ecological circumstances dae (Fig. 2B). Seven families included positive occurrences with a high uncertainty. Variables were continuous and in more than a single genus (see Supplemental Material S4 comprised 17 environmental parameters with a spatial in the Supplemental Materials available online). The three resolution of 30 arc seconds (an approximately 1 km2 grid), families with the most positive occurrences are neither fully which can be classified into three categories: climate, terrestrial nor fully aquatic, as indicated by their mean (6 environment, and altitude (see Supplemental Material S2 standard deviation [SD]) lifetime aquatic indices (Hyper- in the Supplemental Materials available online); for details oliidae, 1.83 6 0.24; Pyxicephalidae, 1.89 6 0.36; and see Penner et al. (2013). In total, 100 ENMs were calculated. Arthroleptidae, 1.6 6 0.39). The genus Hyperolius (Hyper- For each ENM, occurrence records were randomly divided oliidae) had the highest number of positive occurrences and into two groups: 70% (n ¼ 343) for model training and 30% was reported from all six biogeographic regions (Fig. 2B, (n ¼ 147) for model testing, employing subsampling. The Supplemental Material S4A), and had an average lifetime 100 ENMs were averaged into maximum, mean, and aquatic index of 2.00 (Fig. 3; see Supplemental Material S5 minimum predictions. The average of 10 percentile thresh- in the Supplemental Materials available online). The genus 204 Herpetologica 76(2), 2020

FIG. 1.—Batrachochytrium dendrobatidis-positive occurrences detected in African amphibians by date. Positive occurrences are coded by method used to identify Bd: morphology/histology using newly prepared or museum specimens ¼ blue, diagonally hatched; polymerase chain reaction (PCR) or quantitative real-time PCR (qPCR) using samples collected in the field ¼ green; PCR or qPCR from museum specimens ¼ gray, horizontally hatched; genomic sequencing/genotyping ¼ yellow; unknown ¼ black. A color version of this figure is available online.

Amietia (Pyxicephalidae) had the second highest number of northern Africa was predicted to harbor suitable habitat, positive occurrences and was reported from four of six ranging from the Atlantic side of Morocco along the biogeographic regions (Fig. 2B; Supplemental Material Mediterranean coast to Algeria and Tunisia. This area was S4B), and had a mean aquatic index of 2.1 6 0.33 (Fig. 3; previously predicted by Penner et al. (2013) and confirmed Supplemental Material S5). Although we recovered trends of by Moroccan records reported by El Mouden et al. (2011). increasing positive occurrences with higher average lifetime Isolated coastal areas of eastern Libya (Cyrenaica) and aquatic index for both family and genus, neither was Nilotic Egypt are also predicted, but amphibians of these statistically significant (P . 0.05; Fig. 3). areas have not yet been tested for Bd. Large areas in Only 78 positive occurrence records exist for African Ethiopia contain suitable predicted habitat in the Horn of caecilians. Caecilians that have been tested for Bd were Africa, which is consistent with all previous models (Ron collected from Central and East Africa including Cameroon, 2005; Lotters¨ et al. 2009; Rodder¨ et al. 2009; Penner et al. Sa˜o Tome´ and Prıncipe,´ and Tanzania (Fig. 2B inset). Three 2013) and has been confirmed by Gower et al. (2012). families (Dermophiidae, , and Scolecomorphi- Isolated coastal areas of Eritrea (part of the northern region), dae) included Bd records for two genera each while as well as Djibouti and northern Somalia (part of Horn of Indotyphlidae included a single genus, Idiocranium (see Africa), are predicted to be environmentally suitable for Bd, Supplemental Material S6 in the Supplemental Materials but no studies have yet tested samples from these countries. available online). All African caecilians are terrestrial The coastal lowlands of East Africa are generally excluded, (aquatic index ¼ 1; Supplemental Material S5). with ENMs predicting suitable habitats inland at higher altitudes. By contrast, almost the entirety of coastal South Environmental Niche Modeling Africa is predicted while most of Namibia, Botswana, and Overall, the ENMs performed well with a mean (6SD) Zimbabwe are excluded from the models. The ENMs training AUC of 0.950 (0.002) and mean testing AUC of predict numerous areas of suitable habitat for Bd within 0.945 (60.005). All 17 selected variables contributed to the Central Africa. The western side of the Congo Basin is models. The highest contribution came from the lowest modeled as suitable, including the entirety of southern precipitation value (Rain low, 30.6%), followed by the Cameroon, Equatorial Guinea, and Gabon. Large areas in variance in elevation (Elev. Var., 23.9%) and the percentage southern Republic of the Congo and western Democratic of woody vegetation (MODerate Resolution Imaging Spec- Republic of the Congo, as well as smaller regions in coastal troradiometer [MODIS] tree, 9.4%; https://modis.gsfc.nasa. Angola, are also predicted. A number of areas across the gov/data/dataprod/mod44.php). Jackknife testing revealed Congo Basin, which were not previously modeled as suitable the highest value of the maximum temperatures (Temp. in prior studies are identified as such in the present analyses. max high) as the variable with the greatest information The Albertine Rift continues to be a hotspot for Bd with the content when used alone (for details see Supplemental entire region being predicted as highly suitable. Finally, Materials S7–S13 in the Supplemental Materials available suitable habitat is modeled for Bd across West Africa, online). excluding the Dahomey Gap, a nonforested region ranging Results of ENM modeling showed that suitable environ- from eastern Ghana to western Nigeria; however, as stated mental conditions for Bd are found across all seven African by Penner et al. (2013), no positive occurrences have been biogeographic regions (Fig. 4). Only a narrow region in reported west of the Okomu National Park in Nigeria. ZIMKUS ET AL.—CHYTRID IN AFRICAN AMPHIBIANS 205

FIG. 2.—Batrachochytrium dendrobatidis-positive occurrences detected in African amphibians by location and taxonomic group. Positive occurrences are color-coded by biogeographical region for (A) country and region (inset) and (B) family: Anura and (inset). Legend map is modified from Zimkus et al. (2017): (1) Northern Africa, tan; (2) West Africa, dark green; (3) Central Africa, red; (4) Albertine Rift, pink; (5) East Africa, blue; (6) Horn of Africa, light green; and (7) Southern Africa, yellow. 206 Herpetologica 76(2), 2020

FIG. 3.—Positive occurrences with respect to aquatic index. Average lifetime aquatic index ranging from 1 to 3 according to the criteria outlined by Lips et al. (2003) for (A) family and (B) genus. Results were not statistically significant (family, P ¼ 0.29; genus, P ¼ 0.06). A color version of this figure is available online.

Prevalence of Bd in Endangered Species and in Hyla meridionalis and Pelobates varaldii in the Currently 160 African species are listed in categories with Larache-Lakser Lakbir region. One salamander species the highest risk of extinction including extinct in the wild, (Salamandra algira) tested negative for Bd infection by El critically endangered, and endangered (IUCN 2019). We Mouden et al. (2011), but this was prior to the discovery of found reports of Bd infection in 20.62% of these species Bsal as a second pathogenic chytrid fungus infecting (Table 1). A single species listed as extinct in the wild, amphibians (Martel et al. 2013; Spitzen-van der Sluijs et al. Nectophrynoides asperginis (Tanzania), is known to carry Bd 2016; Stegen et al. 2017). infections. Several rare and threatened species of Phrynoba- Horn of Africa.—Only a single known study has been trachus are also known to carry Bd infections and may be conducted in the Horn of Africa, which included a survey of declining (Table 1). For instance, njiomock high-altitude regions (1620–3225 m) on both sides of the Rift was tagged as potentially extinct (currently listed as critically Valley of Ethiopia (Gower et al. 2012; Supplemental Material S1). Batrachochytrium dendrobatidis was detected endangered), as it has not been found since 2010 despite in both the Bale Mountains (eastern side of Rift Valley) and regular surveying of its small range on Mt. Oku, Cameroon Kaffa Region (western side of Rift Valley), with the presence (Doherty-Bone and Gvozdˇ ık´ 2017; IUCN SSC 2019). Similar of Bd detected in 14 species in 10 genera of frogs, although severe declines to virtually zero specimens found since 2010 no dead or dying frogs were found. The overall prevalence are known in the closely related species Phrynobatrachus was 43% among 120 Ethiopian frogs sampled for Bd, with jimzimkusi and P. steindachneri from the Cameroon the highest prevalence detected in Hyperolius (67%) and ˇ ´ Volcanic Line (Doherty-Bone and Gvozdık 2017), although Phrynobatrachus (56%). Gower et al. (2012) noted that a these two have much larger potential ranges (Zimkus and high percentage (40% of more than 60 species) of amphibian Gvozdˇ ık´ 2013). In total, 10 African species with Bd-positive species from this region are endemic, and some are records are listed as critically endangered, including one declining. Although the causes of population decline in , and the population status for each of these is some species of this region are not known, both habitat loss decreasing (Table 1). An additional 22 African species with and chytridiomycosis are suspected to play a role (Gower et Bd-positive records are listed as endangered, with their al. 2012, 2013a). O’Hanlon et al. (2018) have identified the populations decreasing or unknown. All biogeographic zones BdGPL lineage from Ethiopian specimens. except West Africa have at least one endangered species East Africa.—A number of studies have been conducted identified with Bd. Only two of the 32 species listed in in the East African biogeographic region with Bd detected in categories with the highest risk of extinction have had their Kenya, Malawi, Mozambique, and Tanzania (Supplemental Bd lineage identified (Table 1). Material S1). Several studies have investigated the presence of Bd from the western portions of Burundi, Rwanda, and Regional Review Uganda, which are included under the synopsis of the Northern Africa.—El Mouden (2011) reported the first Albertine Rift; however, there is otherwise no information on and only known occurrence of Bd in northern Africa from the presence of the chytrid pathogen in the eastern parts of samples collected from Morocco (Supplemental Material these countries. Kenyan specimens as far back as 1934 have S1). Of the 203 amphibians sampled, representing 6 families tested positive for Bd infection (Vredenburg et al. 2013), and and 9 species from 51 sites, Bd was confirmed in only 4 more recently genotyping has identified the BdGPL lineage amphibians from 3 sites (total prevalence 6%). In addition, in this country (Byrne et al. 2019). A large study in Kenya Bd was found in Discoglossus scovazzi in the Tetouan region along a nearly 800-km transect from the coast to the border ZIMKUS ET AL.—CHYTRID IN AFRICAN AMPHIBIANS 207

FIG. 4.—Map of confirmed records of Bd on the African continent. The three shades of red represent the geographical extent of three different models (light red ¼ maximum; red ¼ mean; dark red ¼ minimum), predicting the potential distribution of Bd. Modeling is based on the conditions of sites with confirmed presence of the pathogen; for niche parameters see Supplemental Material S2 in the Supplemental Materials available online. Positive occurrences are color-coded by method used to identify Bd: morphology/histology (may include museum specimens) ¼ blue squares; polymerase chain reaction (PCR) or quantitative real-time PCR (qPCR) using samples collected in the field ¼ green circles; PCR or qPCR from museum specimens ¼ gray pentagons; genomic sequencing/genotyping ¼ yellow hexagons; unknown ¼ gray triangle. with Uganda investigated the presence of Bd (Kielgast et al. A number of surveys have been conducted in Tanzania 2010). All 11 sites examined were positive for Bd with an with the detection of Bd in both anurans and caecilians. A overall prevalence of 31.5%; Bd was detected in eight survey of tadpoles on Mount Kilimanjaro in northern species, although no dead or symptomatic specimens were Tanzania reported positive Bd infection from species of found. Kielgast et al. (2010) demonstrated an increase in the Amietia (Zancolli et al. 2013). Gower et al. (2013b) reported prevalence of Bd infection with an increase in altitude and a infections in Scolecomorphus and Boulengerula from the decrease in temperature. Tanzanian forests of Nguru, Nguu, and the Uluguru 208 Herpetologica 76(2), 2020

TABLE 1.—Endangered and critically endangered species that have tested positive for Bd. Asterisk indicates that species may be extinct. Red List categories include extinct in the wild (EW), critically endangered (CR), or endangered (EN). Red List categories, criteria, population trends, and estimated occurrence were reported by IUCN (2019). The Cameroonian species Afrixalus lacteus and Hyperolius ademetzi were identified with the lineage BdCAPE by Byrne et al. (2019). N/A indicates not applicable. See Supplemental Material S1, in the Supplemental Materials available online, for more information.

Red List Red List Population Estimated Country Order Family Genus Species category criteria trend occurrence (km2) Cameroon Anura Arthroleptidae Arthroleptis perreti EN B1ab(iii) Unknown 283 Cameroon Anura Arthroleptidae Astylosternus perreti EN B1ab(iii) Decreasing 1717 Cameroon Anura Arthroleptidae Astylosternus ranoides EN B1ab(iii) Decreasing 2452 Cameroon Anura Arthroleptidae Cardioglossa oreas EN B1ab(iii) Unknown 1383 Cameroon Anura Arthroleptidae Leptodactylodon mertensi EN B1ab(iii) Unknown 1426 Cameroon Anura Arthroleptidae Leptodactylodon erythrogaster CR B1ab(iii) Decreasing 34 Cameroon Anura Bufonidae Wolterstorffina mirei EN B1ab(iii) Decreasing 963 Cameroon Anura Hyperoliidae Afrixalus lacteus EN B1ab(iii) Decreasing 1262 Cameroon Anura Hyperoliidae Hyperolius ademetzi EN B1ab(iii) Decreasing 4853 Cameroon Anura Phrynobatrachidae Phrynobatrachus njiomock CR A2ae; Decreasing 14 B1ab(iii)* Cameroon Anura Pipidae Xenopus longipes CR B1ab(iii,v) Decreasing 2 þ2ab(iii,v) Cameroon Gymnophiona Scolecomorphidae Crotaphatrema lamoteii CR B1ab(iii) Decreasing 49 Cameroon, Nigeria border Anura Bufonidae Wolterstorffina parvipalmata CR A3ce Decreasing 50,088 Cameroon, Nigeria border Anura Phrynobatrachidae Phrynobatrachus jimzimkusi CR A2ae Decreasing 10,281 Cameroon, Nigeria border Anura Phrynobatrachidae Phrynobatrachus steindachneri CR A2ae Decreasing 17,177 Democratic Republic Anura Hyperoliidae Chrysobatrachus cupreonitens EN B1ab(iii) Decreasing 541 of the Congo Ethiopia Anura Brevicipitidae Balebreviceps hillmani CR B1ab(iii,v) Decreasing 5 þ2ab(iii,v) Ethiopia Anura Bufonidae Altiphrynoides malcolmi EN B1ab(iii) Decreasing 630 þ2ab(iii) Ethiopia Anura Hyperoliidae Afrixalus clarkei EN B1ab(iii) Unknown 1263 Ethiopia Anura Ericabatrachus baleensis CR B1ab(i,ii,iii) Unknown 1 þ2ab(i,ii,iii) Malawi Anura Pyxicephalidae Amietia johnstoni EN B1ab(iii) Decreasing 663 Malawi Anura Pyxicephalidae Nothophryne broadleyi EN B1ab(iii) Decreasing Not reported Morocco Anura Pelobatidae Pelobates varaldii EN B2ab(iii) Decreasing Not reported Sa˜o Tome´ and Prıncipe´ Anura Hyperoliidae Hyperolius thomensis EN B1ab(iii) Decreasing Not reported Sa˜o Tome´ and Prıncipe´ Anura Ptychadenidae Ptychadena newtoni EN B1ab(iii)þ Decreasing Not reported 2ab(iii) South Africa Anura Arthroleptidae Leptopelis xenodactylus EN B2ab Decreasing 11,112 (ii,iii,iv,v) South Africa Anura Heleophrynidae Heleophryne hewitti EN B1ab(iii)þ Unknown 338 2ab(iii) South Africa Anura Hyperoliidae Hyperolius pickersgilli EN B1ab(ii,iii) Unknown 4768 þ2ab(ii,iii) South Africa Anura Pipidae Xenopus gilli EN B1ab(ii,iii,v) Decreasing 4602 þ2ab(ii,iii,v) South Africa Anura Pyxicephalidae Microbatrachella capensis CR B2ab(ii,iii) Decreasing 1560 Tanzania Anura Bufonidae Nectophrynoides asperginis EW N/A Unspecific N/A Tanzania Anura Hyperoliidae Hyperolius puncticulatus EN B1ab(iii) Decreasing 1625 Tanzania Anura Petropedetidae Arthroleptides yakusini EN B1ab(iii) Unknown 3113

Mountains. Tanzania is also home to Nectophrynoides concluded that chytridiomycosis was the proximate cause of asperginis, Kihansi Spray Toads, which is the only African extinction in the wild of N. asperginis, representing the first amphibian to have become extinct in the wild (Channing et documented case of extinction by disease of an African al. 2009). After the construction of a hydropower dam, the amphibian species. restricted water flow over the falls severely reduced the Few records exist from studies in southern East Africa. A amount of spray in the Kihansi Gorge, leading to a reduction study conducted in the region of Mount Mulanje, an isolated in population numbers in the small habitat where the species inselberg in southern Malawi, reported the presence of Bd in had been found (Channing et al. 2006). Although with eight species (Conradie et al. 2011a). The highest load was mitigation measures the population rebounded, in 2003 the detected in the two species of River Frogs of the genus numbers dropped again, and Bd was one of the implicated Amietia. One additional study detected Bd in an isolated causes (Weldon and du Preez 2004). A captive breeding tadpole of Xenopus muelleri from Lilongwe, Malawi, which program was successful for N. asperginis, but the reintro- was collected in 1969 (Soto-Azat et al. 2010). Despite a duction of the toads was not without issues, including survey of 22 species of frogs and one caecilian from four persistent presence of Bd, threats from adjacent human highland localities in central and northern Mozambique, settlements, lack of technical expertise, and inadequate only a single Bd-positive record exists for Mozambique, with funding (Rija et al. 2011; Makange et al. 2014; Ngalason et al. the pathogen detected in a specimen of Xenopus muelleri 2019). C. Weldon (personal observation) more recently from Mount Riba´ue` (Conradie et al. 2016). There is ZIMKUS ET AL.—CHYTRID IN AFRICAN AMPHIBIANS 209 presently no information on the presence of Bd in Tome´ and Bioko Island specimens (Supplemental Material amphibians in Zambia. S1). Despite this apparent widespread distribution, some Southern Africa.—The presence of Bd in southern surveys in Gabon, Cameroon, and eastern Nigeria (Came- Africa has been demonstrated in 15 different studies for a roon Volcanic Line) have not detected Bd at several sites that diversity of species (Supplemental Material S1). A compre- range from lowland forests (Daversa et al. 2011; Gratwicke et hensive study by Tarrant et al. (2013) mapped Bd occurrence al. 2011) to savannah-forest mosaics (Zimkus and Larson data in South Africa to predict the distribution of the 2013) and montane forests (Doherty-Bone et al. 2008; pathogen across the country. The modeling study showed Blackburn et al. 2010; Reeder et al. 2011; Bala´zˇ et al. 2012). that Bd is found along the wet eastern and coastal regions of Due to the opportunistic nature of these field surveys, most South Africa. Positive records across the region are from a do not include sufficient sample sizes to conclude that range of habitats and environments including very arid negative results indicate low or zero pathogen prevalence at desert and moist, cool, high-altitude mountains. Infections these sites. The amphibians of Angola, Central African have been demonstrated in both threatened species and Republic, Chad, and continental Equatorial Guinea have not those of least concern; South Africa showed an overall yet been surveyed for Bd. prevalence of 26.6% across all species and 14.5% for The Bd pathogen was first detected in caecilians from threatened species. A recent study examined the presence both field-swabbed and pet trade individuals, including of Bd along the Orange River system, which drains much of species from Cameroon (Doherty-Bone et al. 2013; Gower et the country and provides a link between amphibian al. 2013b). Doherty-Bone et al. (2013) reported that 62.4% populations in this region (Verster 2018). A number of of the caecilians from Cameroon tested for Bd were positive. different studies have investigated the lineages of Bd found Gower et al. (2013b) was the first to document mortality in South Africa and have identified the BdGPL, BdCAPE, from chytridiomycosis in a caecilian: a captive Geotrypetes and hybrid lineages in this country (Farrer et al. 2011; seraphini, a species distributed across Central and West Rosenblum et al. 2013; O’Hanlon et al. 2018). Africa, exhibited high genomic equivalents and died during The lack of positive Bd records for many southern African antifungal treatment. Batrachochytrium dendrobatidis was countries is likely due to the low sampling numbers. The also detected with high prevalence in the Sa˜o Tome´ caecilian pathogen has been documented in Botswana, Eswatini (Hydeman et al. 2013, 2017). (Swaziland), Lesotho, Mozambique, and Zimbabwe. The Three studies have made use of retrospective sampling of Okavango Delta of northern Botswana appears to be a likely museum specimens to examine historical pathogen preva- habitat for Bd, but there are scant published studies for this lence in Central Africa. A study of African clawed frogs region. The Kingdom of Eswatini (Swaziland) has only a collected between 1884 and 1994 revealed Bd infections single published record of Bd. The fungus was detected from dating to the 1930s, including a specimen collected in Xenopus muelleri but no specific locality data were reported Cameroon in 1933 (Soto-Azat et al. 2010). A second (Weldon 2005). The eastern and southern escarpments of retrospective study that surveyed a greater diversity of Lesotho bordering the KwaZulu-Natal Province of South amphibians collected in Cameroon between 1908 and 2006 Africa have been partially investigated. Positive Bd records found Bd infections dating to 1934 in lowland habitats but from Lesotho are all from the genus Amietia: A. delalandii, did not detect Bd in historical samples from the Camer- A. vertebralis, and A. hymenopus (Smith et al. 2007; Tarrant oonian highlands (Hirschfeld et al. 2016). Unfortunately, the 2012). Many positive records have been reported in northern sample sizes for historical time periods were far too low to Mozambique (presented as part of East Africa), but the area robustly infer low or zero pathogen prevalence in the south of the Zambezi River has not been investigated, Cameroonian highlands; however, Hirschfeld et al. (2016) although Bd is expected to be present. Only a single study documented disappearances and declines of several high- has been published on Bd in Zimbabwe, which includes a elevation species following the 2008 reports of Bd in the single locality near the southern border (Gericke 2008). Cameroonian highlands, suggesting that Bd may be an Herein we document the first records of Bd in Namibia from emergent pathogen in these montane communities. Finally, specimens of Amietia poyntoni from two sites in Namib- a retrospective study of Bd prevalence in the amphibians of Naukluft National Park. Although the fungus has only been Bioko, Prıncipe,´ and Sa˜oTome´ islands included three documented in central Namibia, further studies will likely specimens collected prior to 1998; one specimen tested show that it is more widespread in the country. positive, indicating that Bd has been on Bioko island since Central Africa.—Batrachochytrium dendrobatidis is 1966 while the earliest records currently known for Prıncipe´ widespread in Central Africa, with recent field surveys of and Sa˜o Tome´ islands date to 2001 (Hydeman et al. 2017). frogs and caecilians reporting the pathogen in Gabon (Bell et Albertine Rift.—Goldberg et al. (2007) first detected Bd al. 2011; Jongsma et al. 2016; Hydeman et al. 2017; this in the Albertine Rift in five amphibian families (Arthrolepti- study, Supplemental Material S3), Cameroon (Doherty- dae, Bufonidae, Hyperoliidae, Ptychadenidae, and Pipidae; Bone et al. 2013; Gower et al. 2013b; Hirschfeld et al. 2016; overall prevalence ¼ 22%) from Kibale National Park in Miller et al. 2018; this study, Supplemental Material S3), the western Uganda in 2006. Greenbaum et al. (2008) conducted oceanic islands of Sa˜o Tome´ and Prıncipe´ (Hydeman et al. surveys in Kahuzi Biega National Park in eastern Democratic 2013, 2017), the land-bridge island Bioko (Hydeman et al. Republic of the Congo 1 yr later, and two specimens/species 2017; P. McLaughlin, personal observation), and the of Hyperolius (Hyperoliidae) were identified to have Bd Republic of the Congo, herein reported for the first time infection via histological identification of sporangia contain- (Supplemental Material S3). O’Hanlon et al. (2018) and ing zoospores in toe clippings. Use of histology and scanning Byrne et al. (2019) have also both recently identified electron microscopy also identified the presence of Bd in BdCAPE in Cameroonian specimens and BdGPL in Sa˜o Ugandan larvae of Amietia ruwenzorica (Pyxicephalidae), a 210 Herpetologica 76(2), 2020 species found in both Uganda and adjacent Democratic Hemisotidae, Hyperoliidae, or Pipidae, and a higher Republic of the Congo (Viertel et al. 2012). prevalence was observed during the dry season (9.23%) Greenbaum et al. (2014) surveyed 20 sites in the when compared to the rainy season (3.06%; P , 0.05). Democratic Republic of the Congo near the border with Penner et al. (2013) did not detect Bd in 793 samples Zambia in 2009 and 2010, comprising 32 species and 56 originating from opportunistic field surveys across West individuals. Frogs from eight of the localities tested positive Africa from Guinea to Benin. Environmental niche modeling for Bd; a total of six families and eight genera were found suggests that appropriate habitat exists in West Africa, and with the fungus. Sample sizes within species were low, with a Penner et al. (2013) hypothesized that the Dahomey Gap, a single specimen for 59% of the species; thus, prevalence dry forest-savanna mosaic, may serve as a distributional could not be reliably estimated. Data from an additional 45 barrier to the spread of the fungus. The only known Bd localities in the eastern Democratic Republic of the Congo record west of the Dahomey Gap is from Morehouse et al. on the western side of the Albertine Rift were analyzed from (2003), who report the presence of Bd from one Xenopus collections made from 2008 to 2011 by Greenbaum et al. tropicalis imported from Ghana. However, the individual (2015), including 7 families, 14 genera, and 66 species. A was tested only after it was imported to the United States, so total of 58 of 166 individuals from 28 sites tested positive for it is unclear when and where within the trade pathway the Bd, resulting in an overall prevalence of 34.9%. individual became infected. It has also been reported that Seimon et al. (2015) integrated data from previous studies infections are highly dependent on the density of individuals in this region by Greenbaum et al. (2008, 2014, 2015) with and, given that pet trade are commonly stored and newly collected data to report on Bd prevalence from 39 sites transported in high densities, the infection may have across the Albertine Rift from swab samples collected originated from another locality or been transmitted by between 2010 and 2014. Of the 17 genera that were sampled another traded species (Schloegel et al. 2009, 2010; Briggs et for the presence of Bd, 7 families and 10 genera had samples al. 2010; Catenazzi et al. 2010). This isolated case could also that were Bd positive, resulting in 19.5% of amphibians have resulted from the use of infected cages previously used testing positive across all sites. Although one Phrynobatra- to export individuals infected with Bd. The positive record of chus was found dead during the surveys and skin swabs were the individual imported from Ghana was, therefore, positive for Bd, histology did not identify chytridiomycosis excluded from ENM analysis by Penner et al. (2013) and and no zoospores were identified, leading the authors to herein. conclude that the animal had a subclinical infection. To understand the history of Bd in the region, Seimon et al. DISCUSSION (2015) also tested an additional 232 museum specimens collected in 1925–1994. A single specimen of Phrynoba- Although the presence of Bd in Africa has long been trachus asper collected in 1950 in the Itombwe highlands of known, recent publications with a global Bd focus do not the Democratic Republic of the Congo was the only mention Africa as an area of particular concern for specimen that tested positive. Soto-Azat et al. (2010) had chytridiomycosis (Scheele et al. 2019). Our ENMs predict previously reported the presence of Bd in two Xenopus laevis a number of Bd hotspots in Africa that are validated by a bunyoniensis (now X. victorianus) museum specimens from large number of positive occurrences in these regions 1934 that were collected from Lake Bunyoni in south- including western Central Africa (e.g., Cameroon, Gabon), western Uganda close to the border with Rwanda. Most South Africa, and the Albertine Rift (e.g., Democratic recently, O’Hanlon et al. (2018) and Byrne et al. (2019) have Republic of the Congo). The addition of 378 new African identified the highly virulent strain BdGPL and hybrid localities where Bd has been detected, including both recent lineages in specimens from Uganda and Burundi, respec- and historic records, has allowed us to further refine tively. Lastly, we report five additional positive occurrences previous continental models of the pathogen’s potential from the Democratic Republic of the Congo (Supplemental distribution (Penner et al. 2013). Our models indicate that Material S3). large areas of the continent include habitat suitable to Bd, West Africa.—Bd was first reported in West Africa from a but many of these regions have not yet been well surveyed, wet mount examination of skin slough recovered from likely due to their remote locations or the political instability Chiromantis rufescens collected from Okomu National Park of particular countries. In particular, Bd is likely to have a in Nigeria (Imasuen et al. 2009), an area situated east of the larger prevalence in the Congo Basin, Ethiopia, and Angola, Dahomey Gap. Complete analyses from 732 frogs from this as well as in North African countries along the Mediterra- protected area sampled over 2007 and 2008 were later nean coastline (i.e., Algeria, Libya, Tunisia). Interestingly, reported based on qPCR examination (Imasuen et al. 2011). records reported in Namibia were predicted both by our Members of 7 families, 8 genera, and 21 species were ENMs herein and by previous modeling (Penner et al. 2013), infected with Bd from Okomu National Park with an indicating that these models may be useful to prioritize infection prevalence of 43.6% for this amphibian communi- regions for Bd surveys. The Upper Guinean Forest region of ty. During the wet season, an overall prevalence of 44% was West Africa (west of Dahomey Gap) continues to have large recorded for infected species while overall infection during areas modeled as suitable habitat for Bd, despite the lack of the dry season was significantly lower (33%; P , 0.05). More any positive records to date (Penner et al. 2013). James et al. recent work by Chinemerem (2017) at the Okomu Rubber (2015) suggests that the identification of cold spots or regions Plantation (north of Okomu National Park) documented Bd where Bd is absent will allow increased understanding into in three families, Arthroleptidae, Dicroglossidae, and Pty- both the biotic and abiotic mechanisms underlying the chadenidae, with an overall prevalence of 5.52%. Batracho- spread of the chytrid pathogen. The lack of documented chytrium dendrobatidis was not detected in the Bufonidae, cases of Bd in West Africa highlights the importance of large- ZIMKUS ET AL.—CHYTRID IN AFRICAN AMPHIBIANS 211 scale investigations in understanding the limits of a 2006). In addition, Bd has been reported from nonamphi- pathogen’s distribution and identifies key areas for surveil- bian hosts including crayfish (McMahon et al. 2013; but see lance to prevent further spread of the pathogen. In Betancourt-Roman et al. 2016). It is unclear whether particular, Ghana contains the most suitable habitat west alternative hosts exist in Africa, however. Although Longcore of the Dahomey Gap and should be closely monitored, et al. (1999) suggested that the fungus may survive though it appears no additional survey results for West Africa saprobically on keratin, a saprobic form of Bd has never have been published since the initial regional study (Penner been observed in the laboratory or in nature. et al. 2013). Likewise, the prediction of suitable habitats in A range of different diagnostic methods have been used to the lowlands of the Congo Basin indicates that more surveys identify Bd in African amphibians including histopathology, in this region are needed to refine our ENMs and better conventional PCR, qPCR, and most recently genome understand the elevational and temperature limits of Bd sequencing or genotyping methods. The histological exam- (Seimon et al. 2015). ination of paraffin-embedded and sectioned skin tissue or toe The analyses herein are the first to examine all known clips was the first approach used to identify Bd zoosporangia reports of Bd in African amphibians in a taxonomic scope. in African amphibians (Weldon et al. 2004; Weldon 2005). Most positive occurrences we report are within the Hyper- The most commonly used method to identify Bd in African oliidae (Hyperolius) and Pyxicephalidae (Amietia). Reed amphibians over the last 15 yr has been qPCR assays using Frogs of the genus Hyperolius have aquatic tadpoles and are swabs collected in the field. This method is considered the primarily arboreal, although they can be encountered on the most reliable and least invasive means of detecting chytrid ground and/or in the leaf litter (Channing and Rodel¨ 2019). infection (Boyle et al. 2004). The use of qPCR over A number of previous studies have reported both medium to traditional PCR allows the quantification of Bd zoospore high prevalence and infection levels in Hyperolius (Kielgast loads, and the use of swabs from newly collected animals et al. 2010; Imasuen et al. 2011; Greenbaum et al. 2014). includes fewer potential sources of cross-contamination River Frogs (Amietia) are large amphibians that are always when compared to museum specimens. It should be noted associated with waterbodies such as permanent streams and that many studies that have included historic specimens have ponds, where they breed and aquatic tadpoles develop attempted to mitigate contamination by rinsing the speci- (Channing and Rodel¨ 2019). Several studies have previously mens with clean 70% ethanol before sampling (Vrendenberg documented Bd infections in Amietia (Kielgast et al. 2010; et al. 2013; Hirschfeld et al. 2016; Hydeman et al. 2017). Conradie et al. 2011b; Viertel et al. 2012; Zancolli et al. 2013; Examination for Bd infection in museum material can be Greenbaum et al. 2014, 2015). Although these species rely completed using histological techniques, but these methods on aquatic habitats for part of their lifecycles, positive Bd are often too destructive for valuable specimens. The use of records for caecilian and species lacking aquatic life- interdental brushes with museum specimens was advocated history stages reinforce the complex ecology of Bd, which over skin swabbing or scraping by Soto-Azat et al. (2009) was once believed to be an exclusively aquatic pathogen. because it did not leave any visible evidence compared to Higher infection prevalence is generally observed in aquatic scraping, and they detected higher counts of genomic species when compared to terrestrial species, and it has been suggested that aggregations at aquatic breeding sites increase equivalents, although they were not statistically significant. the risk of Bd infection (Kriger and Hero 2007). The use of genome or genotyping methods to identify the Previous studies of African amphibians support slightly Bd lineage or lineages is increasingly important because lower prevalence of Bd infection in terrestrial species specific strains have been found to be more virulent than (Conradie et al. 2011a; Gower et al. 2012; Doherty-Bone others (e.g., BdGPL . BdCAPE), with global trade et al. 2013; Greenbaum et al. 2015). Hydeman et al. (2017) facilitating the introduction of lineages into new regions also found that Bd prevalence was lower in terrestrial species (O’Hanlon et al. 2018). Whole-genome sequencing in an than that in riparian species with aquatic larvae but analysis of the global diversity of Bd has been used to comparable to species that are fully aquatic. Interestingly, identify two lineages distributed within Africa: BdGPL and laboratory experiments involving Neotropical species (Bra- BdCAPE; the highly virulent BdGPL has been identified in chycephalus pitanga, Eleutherodactylus johnstonei, Ischnoc- seven countries: Burundi, Kenya, Equatorial Guinea, nema parva) demonstrate that infection loads in direct- Ethiopia, Sa˜oTome´ and Prıncipe,´ South Africa, and Uganda developing species increase rapidly, leading to higher (Supplemental Material S1; O’Hanlon et al. 2018; Byrne et mortality rates relative to species with aquatic larva al. 2019). Two hybrids of BdGPL and BdCAPE have also (Mesquita et al. 2017). Low resistance to Bd in direct been identified in South Africa, demonstrating that Bd is developers may be explained by less exposure to Bd in the continuing to generate novel genomic diversity (O’Hanlon et life cycle of these species, leading to a lack of adaptive al. 2018). In addition, Byrne et al. (2019) discovered an response against the fungus. In addition, Bd spillover from undetermined lineage of Bd from specimens from Burundi, low-load aquatic-breeding frogs can lead to infection and concluding that they may have resulted from coinfection or mortality in direct-developing species (Becker et al. 2019), hybridization. We agree with Seimon et al. (2015) who further underscoring the need to consider terrestrial- suggest that a combination of both molecular and histological breeding species when studying the community-level methods may be most informative; molecular methods can impacts of chytridiomycosis. Furthermore, it is yet unclear be used in surveillance to test for Bd and determine whether how Bd might exist apart from its amphibian host in the endemic or introduced lineages are present while histopa- environment, as there is only a single report of environmen- thology can be used to determine if infection has led to tal swabs testing positive from the site of a mortality event in chytridiomycosis in populations that display higher zoospore Panama (swabs taken from the underside of rocks; Lips et al. loads. 212 Herpetologica 76(2), 2020

Previous global risk assessments identified few African not reported sex of individuals infected and/or sex-specific species with Bd in great danger of extinction (Rodder¨ et al. prevalence rates; therefore, no continental-wide or species- 2009). We have found that many additional imperiled specific assessments regarding this factor can be made at this species are infected with the fungus since this assessment time. over a decade ago, and this increase may be a result of Few African field studies to date were directed exclusively geographically and taxonomically broader Bd testing and at investigating the prevalence and infection intensity of Bd. changes to threat level listings, given decreases in population Most records are by-products of amphibian surveys, and and extent of occurrence. As previously mentioned, the mere much of this work has targeted specific habitats with high occurrence of Bd does not present a threat to species; levels of species richness and endemism, such as high- however, its presence in species at risk should prompt those altitude sky islands; therefore, there is a lack of information involved in conservation programs to determine which about Bd infecting common, widespread lowland species. lineage is present and add Bd surveillance to action plans For example, Seimon et al. (2015) suggested that more if a virulent strain is identified. Batrachochytrium dendro- research be conducted in the lowlands of the Congo Basin batidis is now detected in over one fifth of the African (,600 m) to better refine models and clarify the elevational amphibian species at the greatest risk of extinction, although and temperature limits on Bd infectivity. We advocate that the specific Bd lineage has only been identified in two more basic population-level knowledge is needed for African species to date (Table 1). Phrynobatrachus jimzimkusi amphibians, including studies at smaller spatial and temporal (Critically Endangered) and Cardioglossa oreas (Endan- scales to understand community-level prevalence. In addi- gered) are two species that are herein reported with Bd for tion, more widespread and routine monitoring is needed to the first time, and the dramatic decline of P. jimzimkusi was understand how Bd may be affecting African populations, previously associated with a community-wide increase in Bd whether it is contributing to declines, and possible routes of prevalence (Hirschfeld et al. 2016; IUCN 2019). The only transmission. Doherty-Bone et al. (2019) suggest that documented extinction in the wild of an African amphibian, population monitoring and Bd surveillance of a wide range that of Kihansi Spray Toads (Nectophrynoides asperginis), of African species is urgently needed, given the recent has recently been attributed to chytridiomycosis (C. Weldon, knowledge that Bd is a novel pathogen to the continent, personal observation). This species was likely especially especially since it is unknown which species might be vulnerable to this infectious disease, given its high popula- susceptible to Bd-driven declines. Finally, we suggest that tion density and aggregation behavior, and the epidemic was identifying Bd lineages that infect endangered amphibian likely exacerbated by the environmental conditions that were species should be a priority because the identification of optimal for the fungus. Unfortunately, the IUCN currently either endemic (BdCape) or hypervirulent strains (BdGPL) categorizes 154 African species as Data Deficient, and can inform conservation management efforts and the several recent studies imply that most Data Deficient species potential outcomes of infection. The absence of (visible) might actually qualify for a high-risk category (Tietje and mass mortalities in African amphibians is not necessarily Rodel¨ 2018; Gonza´lez-del-Pliego et al. 2019); thus, it is evidence that disease-related declines have not occurred possible that additional species may be considered threat- (compare Hirschfeld et al. 2016); rather, missing baseline ened once more basic information regarding their population data for understudied regions and populations, and the small size, distribution, life history, and ecology become known. community of active amphibian researchers on this conti- A number of additional factors may influence prevalence nent, make it unlikely that declines would be detected or that of Bd in African amphibians, but data do not currently exist mass die-offs would be observed directly. Consequently, to allow large-scale examinations of these aspects of the fewer documented declines in Africa by previous global disease’s dynamics. Studies in other parts of the world have reviews such as Scheele et al. (2019) may actually reflect that found that Bd loads increase during drought, which is we as a community know less about amphibian populations attributed to the fact that zoospores are concentrated in on this continent relative to other parts of the world. smaller volumes of water with lower flow rates (i.e., dry season), thereby increasing the likelihood of infection Acknowledgments.—We thank all those who provided global position- ing system coordinates or assisted with questions regarding data associated (Piotrowski et al. 2004; Tunstall 2012; Adams et al. 2017). with previously published studies including T. Goldberg, L. Scheinberg, and Correspondingly, studies involving African amphibian com- V. Vrendenburg. Fieldwork in Namibia by AC was completed under Ethics munities support similar patterns regarding seasonal preva- approval UWC 04/4/10; research and collecting permits (unnumbered) were lence, reporting increased prevalence during drier months of obtained from the Ministry for the Environment and Tourism by M. Griffin. the year (Conradie et al. 2011b; Chinemerem 2017), but Fieldwork by EG and CK in Democratic Republic of the Congo was conducted under Permits Nos. LWI/27/BBa/MUH.M/BBY/141/09 (2009) seasonal variation in prevalence is uncharacterized for the and LWI/27/BBa/MUH.M/BBY/001/011 from the Director General of vast majority of the continent. Behavioral factors, such as Centre de Recherche en Sciences Naturelles and was supported by a increased contact among males or increased contact with National Geographic Research and Exploration Grant (No. 8556-08), water, can lead to differences in Bd transmission between University of Texas at El Paso (UTEP), and a grant from the National Science Foundation (NSF; DEB-1145459); field companions Mwenebatu the sexes (Johnson et al. 2003; Carey et al. 2006; Wheeler M. Aristote and Wandege M. Muninga assisted with collecting. Fieldwork in and Welsh 2008). For example, studies have found that Bd the Republic of Congo was completed under NSF research grant No. prevalence is higher in male Rana boylii, a stream-breeding 1145437 to KJ under Groupe d-Etude et de Recherche sur la Diversite´ anuran endemic to the western United States, when Biologique permits No. 108/DIR.08 and No. 037/DIR.10. KJ would like to compared to females or juveniles (Adams et al. 2017). Some thank the Smithsonian National Museum of Natural History (USNM) Herpetology Department, the Wildlife Conservation Society (WCS) Congo, studies on African amphibian populations have reported that and Institut National de Recherche en Sciences Exactes et Naturelles Bd prevalence was higher in males when compared to (IRSEN) in Brazzaville. Fieldwork by JGL in Gabon was completed under females (Imasuen 2012; Chinemerem 2017), but most have Parcs Gabon Autorisation d’Entree dans les Parcs Nationaux No. AE130014/ ZIMKUS ET AL.—CHYTRID IN AFRICAN AMPHIBIANS 213

PR/ANPN/SE/CS/AEPN, Autorisation de Recherches sur le Territoire du Catenazzi, A., V.T. Vredenburg, and E. Lehr. 2010. Batrachochytrium Gabon No. AR0013/13/MENESTFPRSCJS/CENAREST/CG/CST/CSAR, dendrobatidis in the live frog trade of Telmatobius (Anura: Cerato- and CERTIFICAT D’ORIGINE No 79-84/DGFAP POUR LES PRO- phryidae) in the tropical Andes. Diseases of Aquatic Organisms 92:187– DUITS EXPORTES. Funding for labwork on Gabonese samples was 191. provided by the University of Michigan to JGL and TYJ. Fieldwork by VG in Channing, A., and M.-O. Rodel.¨ 2019. Field Guide to the Frogs and Other Cameroon was completed under Permits 1010/PRBS/MINFOF/SG/DFAP/ Amphibians of Africa. Struik Nature, South Africa. SDVEF/SC and 1099/PRS/MINFOF/SG/DFAP/SDVEF/SC. VG would like Channing, A., K.S. Finlow-Bates, S.E. Haarklau, and P.G. Hawkes. 2006. to thank M. Dolinay and G.C. Tasse Taboue for help in the field and R. The biology and recent history of the critically endangered Kihansi Spray Tropek and A. Koda´dkova´ for additional material. VG’s research was Toad Nectophrynoides asperginis in Tanzania. Journal of East African supported by the Czech Science Foundation, Grant Number 15-13415Y, Natural History 95:117–138. and the Ministry of Culture of the Czech Republic (DKRVO 2019–2023/ Channing, A., K. Howell, S. Loader, M. Menegon, and J. Poynton. 2009. 6.VII.b, National Museum, 00023272). BZ is grateful to E. Ballerini and M. Nectophrynoides asperginis. In IUCN 2010: IUCN Red List of Laslo for assistance with RStudio and to D. Green for the invitation to speak Threatened Species, Version 2010.1. Available at http://www. at the amphibian population declines symposium that initiated this work. iucnredlist.org. Accessed on 7 January 2020. International Union for Conservation of Nature, Switzerland. SUPPLEMENTAL MATERIAL Chinemerem, I.G. 2017. A Survey of Chytridiomycosis among Amphibians (Anurans) in Okomu Rubber Plantation, Edo State Nigeria. M.S. thesis, Supplemental material associated with the article can be University of Benin, Nigeria. found online at http://dx.doi.org/10.1655/ Conradie, W., J. Harvey, A. Kotze,´ D.L. Dalton, and M.J. Cunningham. HERPETOLOGICA-D-19-00058.S1 and via the Museum 2011a. Confirmed amphibian chytrid in Mount Mulanje Area, Malawi. f¨ur Naturkunde Berlin (MfN) at https://doi.org/10.7479/ Herpetological Review 42:369–371. enkp-cb58. Conradie, W., C. Weldon, K.G. Smith, and L.H. du Preez. 2011b. Seasonal pattern of chytridiomycosis in common river frog (Amietia angolensis) tadpoles in the South African Grassland Biome. African Zoology 46:95– 102. ITERATURE ITED L C Conradie, W., G.B. Bittencourt-Silva, S.P. Loader, D.L. Dalton, and K.A. Adams, A.J., S.J. Kupferberg, M.Q. Wilber, A.P. Pessier, M. Grefsrud, S. Tolley. 2016. Batrachochytrium dendrobatidis survey of amphibians in Bobzien, V.T. Vredenburg, and C.J. Briggs. 2017. Extreme drought, host the northern Mozambique ‘‘Sky Islands’’ and low-lying areas. Herpeto- density, sex, and bullfrogs influence fungal pathogen infection in a logical Review 47:42–46. declining lotic amphibian. Ecosphere 8:e01740. Daversa, D., J. Bosch, and K. Jeffery. 2011. First survey of the chytrid Bala´z,ˇ V., O. Kopecky,´ and V. Gvozdˇ ık.´ 2012. Presence of the amphibian fungus, Batrachochytrium dendrobatidis, in amphibian populations of chytrid pathogen confirmed in Cameroon. Herpetological Journal Gabon, Africa. Herpetological Review 42:67–69. 22:191–194. Doherty-Bone, T.M., and V. Gvozdˇ ık.´ 2017. The amphibians of Mount Oku, Becker, C.G., M.C Bletz, S.E. Greenspan,. . . C.F.B. Haddad. 2019. Low- Cameroon: An updated species inventory and conservation review. load pathogen spillover predicts shifts in skin microbiome and survival of ZooKeys 643:109–139. a terrestrial-breeding amphibian. Proceedings of the Royal Society B Doherty-Bone, T.M., J. Bielby, N.L. Gonwouo, M. LeBreton, and A.A. 286:20191114. Cunningham. 2008. In a vulnerable position? Preliminary survey work Bell, R.C., A.V. Gata Garcia, B.L. Stuart, and K.R. Zamudio. 2011. High fails to detect the amphibian chytrid pathogen in the highlands of prevalence of the amphibian chytrid pathogen in Gabon. EcoHealth Cameroon, an amphibian hotspot. Herpetological Journal 18:115–118. 8:116–120. Doherty-Bone, T.M., N.L. Gonwouo, M. Hirschfeld,. . . A.A. Cunningham. Bell, R.C., R.C. Drewes, A. Channing, V. Gvozdˇ ık,´ J. Kielgast, S. Lotters,¨ 2013. Batrachochytrium dendrobatidis in amphibians of Cameroon, B.L. Stuart, and K.R. Zamudio. 2015. Overseas dispersal of Hyperolius including first records of infected caecilian hosts. Diseases of Aquatic ˜ ´ reed frogs from Central Africa to the oceanic islands of Sao Tome and Organisms 102:187–194. Prıncipe.´ Journal of Biogeography 42:65–75. Doherty-Bone, T.M., A.A. Cunningham, M.C. Fisher, T.W.J. Garner, P. Berger, L., R. Speare, P. Daszak,. . . A.A. Cunningham. 1998. Chytridiomy- Ghosh, D.J. Gower, R. Verter, and C. Weldon. 2019. Amphibian chytrid cosis causes amphibian mortality associated with population declines in fungus in Africa—Realigning hypotheses and the research paradigm. the rain forests of Australia and Central America. Proceedings of the Animal Conservation 2019:1–6. DOI: http://dx.doi.org/10.1111/acv. National Academy of Sciences of the United States of America 95:9031– 12538. 9036. El Mouden, E.H., T. Slimani, D. Donaire, S. Ferna´ndez-Beaskoetxea, M.C. Berger, L., A.D. Hyatt, R. Speare, and J.E. Longcore. 2005. Life cycle stages of Batrachochytrium dendrobatidis. Diseases of Aquatic Organisms Fisher, and J. Bosch. 2011. First record of the chytrid fungus 68:51–63. Batrachochytrium dendrobatidis in North Africa. Herpetological Review Betancourt-Roman, C.M., C.C. O’Neil, and T.Y. James. 2016. Rethinking 42:71–75. the role of invertebrate hosts in the life cycle of the amphibian Elith, J., C.H. Graham, R.P. Anderson,. . .N.E. Zimmermann. 2006. Novel chytridiomycosis pathogen. Parasitology 143:1723–1729. methods improve prediction of species’ distributions from occurrence Blackburn, D.C., B.J. Evans, A.P. Pessier, and V.T. Vredenburg. 2010. An data. Ecography 29:129–151. enigmatic mortality event in the only population of the critically Elith, J., S.J. Phillips, T. Hastie, M. Dudık,´ Y. Chee, and C.J. Yates. 2011. A endangered Cameroonian frog Xenopus longipes. African Journal of statistical explanation of MaxEnt for ecologists. Diversity and Distribu- Herpetology 59:111–122. tions 17:43–57. Boyle, D.G., D.B. Boyle, V. Olsen, J.A. Morgan, and A.D. Hyatt 2004. Rapid Farrer, R.A., L.A. Weinert, J. Bielby,. . . M.C. Fisher. 2011. Multiple quantitative detection of chytridiomycosis (Batrachochytrium dendroba- emergences of genetically diverse amphibian-infecting chytrids include a tidis) in amphibian samples using real-time Taqman PCR assay. Diseases globalized hypervirulent recombinant lineage. Proceedings of the of Aquatic Organisms 60:141–148. National Academy of Sciences of the United States of America Briggs, C.J., R.A. Knapp, and V.T. Vredenburg. 2010. Enzootic and 108:18732–18736. epizootic dynamics of the chytrid fungal pathogen of amphibians. Frost, D.R. 2020. Amphibian Species of the World: An Online Reference, Proceedings of the National Academy of Sciences of the United States Version 6.0. Available at http://research.amnh.org/herpetology/amphibia/ of America 107:9695–9700. index.html. Accessed on 7 January 2020. American Museum of Natural Byrne, A.Q., V.T. Vredenburg, A. Martel,. . . E.B. Rosenblum. 2019. Cryptic History, USA. diversity of a widespread global pathogen reveals new threats for Gericke, M.C. 2008. Aspects of Amphibian Chytrid Infections in South amphibian conservation. Proceedings of the National Academy of Africa. M.S. thesis, North-West University, Potchefstroom, South Africa. Sciences of the United States of America 116:20382–20387. Goldberg, T.L., A.M. Readel, and M.H. Lee. 2007. Chytrid fungus in frogs Carey, C., J.E. Bruzgul, L.J. Livo, M.L. Walling, K.A. Kuehl, B.F. Dixon, from an equatorial African montane forest in western Uganda. Journal of A.P. Pessier, R.A. Alford, and K.B. Rogers. 2006. Experimental exposures Wildlife Diseases 43:521–524. of boreal toads (Bufo boreas) to a pathogenic chytrid fungus (Batracho- Gonza´lez-del-Pliego, P., R.P. Freckleton, D.P. Edwards, M.S. Koo, B.R. chytrium dendrobatidis). EcoHealth 3:5–21. Scheffers, R.A. Pyron, and W. Jetz. 2019. Phylogenetic and trait-based 214 Herpetologica 76(2), 2020

prediction of extinction risk for data-deficient amphibians. Current occurrence of the amphibian chytrid fungus in Kenya. Animal Biology 29:1557–1563. Conservation 13:1–8. Gower, D.J., T.M. Doherty-Bone, R.K. Aberra, A. Mengistu, S. Schwaller, Kriger, K.M., and J.-M Hero. 2007. The chytrid fungus Batrachochytrium M. Menegon, R.O. de Sa´, S.A. Saber, A.A. Cunningham, and S.P. Loader. dendrobatidis is non-randomly distributed across amphibian breeding 2012. High prevalence of the amphibian chytrid fungus (Batrachochy- habitats. Diversity and Distributions 13:781–788. trium dendrobatidis) across multiple taxa and localities in the highlands Leache,´ A.D., P. Wagner, C.W. Linkem,. . . F.T. Burbrink. 2014. A hybrid of Ethiopia. Herpetological Journal 22:225–233. phylogenetic–phylogenomic approach for species tree estimation in Gower, D.J., R.K. Aberra, S. Schwaller,. . . S.P. Loader. 2013a. Long-term African Agama lizards with applications to biogeography, character data for endemic frog genera reveal potential conservation crisis in the evolution, and diversification. Molecular Phylogenetics and Evolution Bale Mountains, Ethiopia. Oryx 47:59–69. 79:215–230. Gower, D.J., T. Doherty-Bone, S.P. Loader,. . . T.W.J. Garner. 2013b. Linder, H.P., H.M. de Klerk, J. Born, N.D. Burgess, J. Fjeldsa,˚ and C. Batrachochytrium dendrobatidis infection and lethal chytridiomycosis in Rahbek. 2012. The partitioning of Africa: Statistically defined biogeo- caecilian amphibians (Gymnophiona). EcoHealth 10:173–183. graphical regions in sub-Saharan Africa. Journal of Biogeography Gratwicke, B., A. Alonso, T. Elie, J. Kolowski, J. Lock, N. Rotzel, J. Sevin, 39:1189–1205. and R.C. Fleischer. 2011. Batrachochytrium dendrobatidis not detected Lips, K.R., J.D. Reeve, and L.R. Witters. 2003. Ecological traits predicting on amphibians from two lowland sites in Gabon, Africa. Herpetological amphibian population declines in Central America. Conservation Biology Review 42:69. 17:1078–1088. Greenbaum, E., C. Kusamba, M.M. Aristote, and K. Reed. 2008. Amphibian Lips, K.R., F. Brem, R. Brenes, J.D. Reeve, R.A. Alford, J. Voyles, C. Carey, chytrid fungus infections in Hyperolius (Anura: Hyperoliidae) from L. Livo, A.P. Pessier, and J.C. Collins. 2006. Emerging infectious disease eastern Democratic Republic of Congo. Herpetological Review 39:70–73. and the loss of biodiversity in a Neotropical amphibian community. Greenbaum, E., J. Meece, K. Reed, and C. Kusamba. 2014. Amphibian Proceedings of the National Academy of Sciences of the United States of chytrid infections in non-forested habitats of Katanga, Democratic America 103:3165–3170. Republic of the Congo. Herpetological Review 45:610–614. Longcore, J.E., A.P. Pessier, and D.K. Nichols. 1999. Batrachochytrium Greenbaum, E., J. Meece, K.D. Reed, and C. Kusamba. 2015. Extensive dendrobatidis gen. et sp. nov., a chytrid pathogenic to amphibians. occurrence of the amphibian chytrid fungus in the Albertine Rift, a Mycologia 91:219–227. Central African amphibian hotspot. Herpetological Journal 25:91–100. Lotters,¨ S., J. Kielgast, J. Bielby, S. Schmidtlein, J. Bosch, M. Veith, S. Gurdon, J.B., and N. Hopwood. 2000. The introduction of Xenopus laevis Walker, M.C. Fisher, and D. Rodder.¨ 2009. The link between rapid into developmental biology: Of empire, pregnancy testing and ribosomal enigmatic amphibian decline and the globally emerging chytrid fungus. genes. International Journal of Developmental Biology 44:43–50. EcoHealth 6:358–372. Hirschfeld, M., D.C. Blackburn, T.M. Doherty-Bone, L.N. Gonwouo, S. Makange, M., N. Kulaya, E. Biseko, P. Kalenga, S, Mutagwaba and G. Ghose, and M-O. Rodel.¨ 2016. Dramatic declines of montane frogs in a Misinzo. 2014. Batrachochytrium dendrobatidis detected in Kihansi Central African biodiversity hotspot. PLoS One 11:e0155129. spray toads at a captive breeding facility (Kihansi, Tanzania). Diseases of Hydeman, M.E., R.C. Bell, R.C. Drewes, and K.R. Zamudio. 2013. Aquatic Organisms 111:159–164. Amphibian chytrid fungus confirmed in endemic frogs and caecilians Martel, A., A. Spitzen-van der Sluijs, M. Blooi,. . . F. Pasmans. 2013. on the Island of Sa˜o Tome,´ Africa. Herpetological Review 44:254–257. Batrachochytrium salamandrivorans sp. nov. causes lethal chytridiomy- Hydeman, M.E., A.V. Longo, G. Velo-Anton,´ D. Rodriguez, K.R. Zamudio, cosis in amphibians. Proceedings of the National Academy of Sciences of and R.C. Bell. 2017. Prevalence and genetic diversity of Batrachochy- the United States of America 110:15325–153259. trium dendrobatidis in Central African island and continental amphibian Martel, A., M. Vila-Escale, D. Ferna´ndez-Giberteau,. . . F. Pasmans. 2020. communities. Ecology and Evolution 7:7729–7738. Integral chain management of wildlife diseases. Conservation Letters Imasuen, A.A. 2012. Investigation of the Helminth Parasitic Infections and 2020:e12707. DOI: https://dx.doi.org/10.1111/conl.12707. Chytridiomycosis of Amphibians in Okomu National Park, Nigeria. Ph.D. McMahon, T.A., L.A. Brannelly, M.W.H. Chatfield, P.T.J. Johnson, M.B. thesis, University of Benin, Nigeria. Joseph, V.J. McKenzie, C.L. Richards-Zawacki, M.D. Venesky, and J. R. Imasuen, A.A., C. Weldon, M.S.O. Aisien, and L. H. du Preez. 2009. Rohr. 2013. Chytrid fungus Batrachochytrium dendrobatidis has non- Amphibian chytridiomycosis: First report in Nigeria from the skin of amphibian hosts and releases chemicals that cause pathology in the Chiromantis rufescens. Froglog 90:6–8. absence of infection. Proceedings of the National Academy of Sciences of Imasuen, A.A., M.S.O. Aisen, C. Weldon, D.L. Dalton, A. Kotze, and LH. the United States of America 110:210–215. du Preez. 2011. Occurrence of Batrachochytrium dendrobatidis in Measey, G.J., M. Vences, R.C. Drewes, Y. Chiari, M. Melo, and B. Bourles. amphibian populations of Okomu National Park, Nigeria. Herpetological 2007. Freshwater paths across the ocean: Molecular phylogeny of the frog Review 42:379–382. Ptychadena newtoni gives insights into amphibian colonization of oceanic IUCN (International Union for Conservation of Nature). 2019. The IUCN islands. Journal of Biogeography 34:7–20. Red List of Threatened Species, Version 2019-2. Available at http://www. Mesquita, A.F.C., C. Lambertini, M. Lyra, L.R. Malagoli, T.Y. James, L.F. iucnredlist.org. Accessed on 10 August 2019. International Union for Toledo, C.F.B. Haddad, and C.G. Becker. 2017. Low resistance to Conservation of Nature, Switzerland. chytridiomycosis in direct developing amphibians. Science Reports IUCN SSC (International Union for Conservation of Nature Species 7:16605:1–7. Survival Commission, Amphibian Specialist Group). 2019. Phrynoba- Miller, C.A., G.C. Tasse Taboue, M.M.P. Ekane, M. Robak, P.R. Sesink trachus njiomock. The IUCN Red List of Threatened Species Clee, C. Richards-Zawacki, E.B. Fokam, N.A. Fuashi, and N.M. 2019:e.T76317553A76317920. Available at http://dx.doi.org/10.2305/ Anthony. 2018. Distribution modeling and lineage diversity of the chytrid IUCN.UK.2019-1.RLTS.T76317553A76317920.en. Accessed on 15 Au- fungus Batrachochytrium dendrobatidis (Bd) in a central African gust 2019. International Union for Conservation of Nature, Switzerland. amphibian hotspot. PLoS One 13:e0199288. James, T.Y., L.F. Toledo, D. Rodder,.¨ . . J.E. Longcore. 2015. Disentangling Morehouse, E.A, T.Y. James, A.R.D. Ganley, R. Vilgalys, L. Berger, P.J. host, pathogen, and environmental determinants of a recently emerged Murphy, and J.E. Longcore. 2003. Multilocus sequence typing suggests wildlife disease: Lessons from the first 15 years of amphibian that the chytrid pathogen of amphibians is a recently emerged clone. chytridiomycosis research. Ecology and Evolution 5:4079–4097. Molecular Ecology 12:395–403. Jongsma, G.F.M., A.B. Kaya, J.-A. Yoga,. . . D.C. Blackburn. 2016. Ngalason, W., C.L. Nahonyo, and C.A. Msuya. 2019. The dynamics of re- Widespread presence and high prevalence of Batrachochytrium den- introduced Kihansi Spray Toad Nectophrynoides asperginis and other drobatidis in Gabon. Herpetological Review 47:227–230. amphibians in Kihansi Gorge, Udzungwa Mountains, Tanzania. Tanzania Johnson, M.L., L. Berger, L. Phillips, and R. Speare. 2003. Fungicidal Journal of Science 45:584–598. effects of chemical disinfectants, UV light, desiccation and heat on the O’Hanlon, S.J., A. Rieux, R.A., Farrer,. . . M.C. Fisher. 2018. Recent Asian amphibian chytrid, Batrachochytrium dendrobatidis. Diseases of Aquatic origin of chytrid fungi causing global amphibian declines. Science Organisms 57:255–260. 627:621–627. Johnson, P.T.J. 2006. Amphibian diversity: Decimation by disease. Penner, J., M. Wegmann, A. Hillers, M. Schmidt, and M.-O. Rodel.¨ 2011. A Proceedings of the National Academy of Sciences of the United States hotspot revisited—A biogeographical analysis of West African amphib- of America 103:3011–3012. ians. Diversity and Distributions 17:1077–1088. Kielgast, J., D. Rodder,¨ M. Veith, and S. Lotters.¨ 2010. Widespread Penner, J., G.B. Adum, M.T. McElroy,. . . M.-O. Rodel.¨ 2013. West Africa— ZIMKUS ET AL.—CHYTRID IN AFRICAN AMPHIBIANS 215

A safe haven for frogs? A sub-continental assessment of the chytrid Stegen, G., F. Pasmans, B.R. Schmidt,. . . A. Martel. 2017. Drivers of fungus (Batrachochytrium dendrobatidis). PLoS One 8:e56236. salamander extirpation mediated by Batrachochytrium salamandrivor- Peterson, A.T., J. Soberon,´ R.G. Pearson, R.P. Anderson, E. Martınez-´ ans. Nature 544:353–356. Meyer, M. Nakamura, and M.B. Araujo.´ 2011. Ecological Niches and Stuart, S.N., J.S. Chanson, N.A. Cox, B.E. Young, A.S. Rodrigues, D.L. Geographic Distributions (MPB-49). Princeton University Press, UK and Fischman, and R.W. Waller. 2004. Status and trends of amphibian USA. declines and extinctions worldwide. Science 306:1783–1786. Piotrowski, J., S. Annis, and J. Longcore. 2004. Physiology of Batrachochy- Tarrant, J. 2012. Conservation Assessment of Threatened Amphibians in trium dendrobatidis, a chytrid pathogen of amphibians. Mycologia 96:9– KwaZulu-Natal, and a National Assessment of Chytrid Infection in South 15. African Threatened Species. Ph.D. dissertation, North-West University, Phillips, S.J., M. Dudık,´ and RE. Schapire. 2019. maxent software for Potchefstroom, South Africa. modeling species niches and distributions, Version 3.4.1. Available at Tarrant, J., D. Cilliers, L.H. du Preez, and C. Weldon. 2013. Spatial http://biodiversityinformatics.amnh.org/open_source/maxent/. Accessed assessment of amphibian chytrid fungus (Batrachochytrium dendrobati- on 10 August 2019. dis) in South Africa confirms endemic and widespread infection. PLoS Qiao, H., J. Soberon,´ and A.T. Peterson. 2015. No silver bullets in correlative One 8:e69591. ecological niche modelling: Insights from testing among many potential Tietje, M., and M.-O. Rodel.¨ 2018. Evaluating the predicted extinction risk algorithms for niche estimation. Methods in Ecology and Evolution of living amphibian species with the fossil record. Ecology Letters 6:1126–1136. 21:1135–1142. R Core Team. 2019. R: A language and environment for statistical Tunstall, T.S. 2012. Characteristics of the Emergent Disease Batrachochy- computing, RStudio Version 1.2.1335. Available at https://www.R- project.org/. R Foundation for Statistical Computing, Austria. trium dendrobatidis in the Rana muscosa and Rana sierrae Species Reeder, N.M.M., T.L. Cheng, V.T. Vredenburg, and D.C. Blackburn. 2011. Complex. Ph.D. dissertation, University of California, USA. Survey of the chytrid fungus Batrachochytrium dendrobatidis from Verster, J.R. 2018. Spatial Epidemiology of Amphibian Chytridiomycosis in montane and lowland frogs in eastern Nigeria. Herpetology Notes 4:83– the Orange River System of South Africa. M.S. thesis, North-West 86. University, South Africa. Rija, A.A., F.H. Khatibu, E.M. Kohi, and R. Muheto. 2011. Status and Viertel, B., M. Veith, S. Schick, A. Channing, S. Kigoolo, O. Baeza-Urrea, U. reintroduction of the Kihansi Spray Toad Nectophrynoides asperginis in Sinsch, and S. Lotters.¨ 2012. The stream-dwelling larva of the Ruwenzori Kihansi Gorge: Challenges and opportunities. Pp. 11–20 in Proceedings river frog, Amietia ruwenzorica, its buccal cavity and pathology of of the 7th TAWIRI Scientific Conference. Tanzania Wildlife Research chytridiomycosis. Zootaxa 3400:43–57. Institute, Tanzania. Vredenburg, V.T., S.A. Felt, E.C. Morgan, S.V.G. McNally, S. Wilson, and Rodder,¨ D., J. Kielgast, J. Bielby, S. Schmidtlein, J. Bosch, T.W.J. Garner, S.L. Green. 2013. Prevalence of Batrachochytrium dendrobatidis in M. Veith, S. Walker, M.C. Fisher, and S. Lotters.¨ 2009. Global amphibian Xenopus collected in Africa (1871–2000) and in California (2001–2010). extinction risk assessment for the panzootic chytrid fungus. Diversity PLoS One 8:e63791. 1:52–66. Wake, D.B., and V.T. Vredenburg. 2008. Are we in the midst of the sixth Ron, S.R. 2005. Predicting the distribution of the amphibian pathogen mass extinction? A view from the world of amphibians. Proceedings of Batrachochytrium dendrobatidis in the New World. Biotropica 37:209– the National Academy of Sciences of the United States of America 221. 105:11466–11473. Rosenblum, E.B., T.Y. James, K.R. Zamudio,. . . J.E. Stajich. 2013. Complex Weldon, C. 2005. Chytridiomycosis, an Emerging Infectious Disease of history of the amphibian-killing chytrid fungus revealed with genome Amphibians in South Africa. Ph.D. dissertation, North-West University, resequencing data. Proceedings of the National Academy of Sciences of South Africa. the United States of America 110:9385–9390. Weldon, C., and L. du Preez. 2004. Decline of the Kihansi spray toad, Scheele, B., F. Pasmans, L. Skerratt,. . . S. Canessa. 2019. Amphibian fungal Nectophrynoides asperginis, from the Udzungwa mountains, Tanzania. panzootic causes catastrophic and ongoing loss of biodiversity. Science Froglog 62:2–3. 363:1459–1463. Weldon, C., L.H. du Preez, A.D. Hyatt, R. Muller, and R Speare. 2004. Schloegel, L.M., A.M. Picco, A.M. Kilpatrick, A.J. Davies, A.D. Hyatt, and Origin of the amphibian chytrid fungus. Emerging Infectious Diseases P. Daszak. 2009. Magnitude of the US trade in amphibians and presence 10:2100–2105. of Batrachochytrium dendrobatidis and ranavirus infection in imported Weldon, C., L. de Villiers, and L.H. du Preez. 2007. Quantification of the North American bullfrogs (Rana catesbeiana). Biological Conservation trade in Xenopus laevis from South Africa, with implications for 142:1420–1426. biodiversity conservation. African Journal of Herpetology 56:77–83. Schloegel, L.M., C.M. Ferreira, T.Y. James,. . . P. Daszak. 2010. The North Wheeler, C.A., and H.H. Welsh. 2008. Mating strategy and breeding American bullfrog as a reservoir for the spread of Batrachochytrium patterns of the foothill yellow-legged frog (Rana boylii). Herpetological dendrobatidis in Brazil. Animal Conservation 13:53–61. Conservation and Biology 3:128–142. Seimon, T.A., S. Ayebare, R. Sekisambu,. . . A.J. Plumptre. 2015. Assessing the threat of amphibian chytrid fungus in the Albertine Rift: Past, present Wickham, H. 2016. ggplot2: Elegant Graphics for Data Analysis. Springer- and future. PLoS One 10:e0145841. Verlag, USA. Skerratt, L., L. Berger, R. Speare, S. Cashins, K.R. McDonald, A.D. Phillott, Zancolli, G., A. Storfer, and M.-O. Rodel.¨ 2013. Detection of Batrachochy- H.B. Hines, and N. Kenyon. 2007. Spread of chytridiomycosis has caused trium dendrobatidis in river frogs (genus Amietia) on Mount Kilimanjaro, the rapid global decline and extinction of frogs. EcoHealth 4:125–134. Tanzania. Herpetological Review 44:611–614. Smith, K.G., C. Weldon, W. Conradie, and L.H. du Preez. 2007. Zimkus, B.M., and V. Gvozdˇ ık.´ 2013. Sky Islands of the Cameroon Volcanic Relationships among size, development, and Batrachochytrium dendro- Line: A diversification hotspot for puddle frogs (Phrynobatrachidae: batidis infection in African tadpoles. Diseases of Aquatic Organisms Phrynobatrachus). Zoologica Scripta 42:591–611. 74:159–164. Zimkus, B.M., and J.G. Larson. 2013. Assessment of the amphibians of Soto-Azat., C., B.T. Clarke, M.C. Fisher, S.F. Walker, and A.A. Cunning- Batek´ e´ Plateau National Park, Gabon, including results of chytrid ham. 2009. Non-invasive sampling methods for the detection of pathogen tests. Salamandra 49:159–170. Batrachochytrium dendrobatidis in archived amphibians. Diseases of Zimkus, B.M., L.P. Lawson, M.F. Barej,. . . S. Lotters.¨ 2017. Leapfrogging Aquatic Organisms 84:163–166. into new territory: How Mascarene ridged frogs diversified across Africa Soto-Azat, C., B.T. Clarke, J.C. Poynton, and A.A. Cunningham. 2010. and Madagascar to maintain their ecological niche. Molecular Phyloge- Widespread historical presence of Batrachochytridium dendrobatidis in netics and Evolution 106:254–269. African pipid frogs. Diversity and Distributions 16:126–131. Spitzen-van der Sluijs, A., A. Martel, J. Asselberghs,. . . S. Lotters.¨ 2016. Expanding distribution of lethal amphibian fungus Batrachochytrium salamandrivorans in Europe. Emerging Infectious Diseases 22:1286– Accepted on 8 February 2020 1288. Associate Editor: David Lesbarreres` SUPPLEMENT

MATERIALS AND METHODS

Sampling Techniques.— Field expeditions were conducted in Cameroon (2009, 2016; the most recent including sampling at the Cameroon-Nigeria border), the Democratic Republic of the Congo (2009, 2010, 2013), Gabon (2013), Namibia (2007), and the Republic of the Congo (2008, 2010). Various techniques were used to collect frog specimens and samples to test for presence of Bd, which are outlined below, including the collection of toe tips, swab samples in the field, and use of interdental brushes on museum specimens. Cameroonian samples collected in 2016 were collected as standard swabs from live specimens. All frogs were caught by hand using latex gloves or polypropylene bags during night encounter surveys. Swab samples were taken by firmly running sterile cotton swabs (Dryswab MW100 finetip) over the ventral surface, flanks and feet in the standardized manner (Hyatt et al., 2007). Frogs from Cameroon (2009, 2016) were preserved in 75% ethanol as museum voucher specimens and were deposited in the herpetology collection of the National Museum in Prague (NMP), Czech Republic; some specimens are currently in the personal collection of VG at the Institute of Vertebrate Biology of the Czech Academy of Sciences but will be deposited in the NMP in the future. Skin samples from specimens originating from the 2009 Cameroonian trips were collected by interdental brushes in 2011. Frogs from the Democratic Republic of the Congo (2009, 2010, 2013), as well as those from the Republic of the Congo (2008, 2010), were collected by hand (without gloves) or dipnet following audio recording of male advertisement calls (in the case of calling males), or during opportunistic visual encounters. To assist with specimen organization and subsequent photographing, frogs were deposited into well-ventilated and labeled plastic containers immediately after capture. Frogs were usually captured individually, but rarely, individuals of the same species (up to six individuals) were kept in the same container. Plastic containers were disinfected with a 10% bleach solution after each collection event. A subset of individuals, mostly post-metamorphic, were swabbed in their containers for approximately 20 seconds on the ventral surfaces of the abdomen, throat, hands and feet. Frogs were swabbed with sterile swabs (MW100; Medical Wire & Equipment Co., Crosham, UK), which were stored individually and dry in shaded containers away from direct sunlight and heat. Swabs were stored at 4°C upon arrival to the laboratory until laboratory work was completed. Voucher specimens from the Democratic Republic of the Congo were deposited in the University of Texas at El Paso Biodiversity Collections, while those from the Republic of the Congo were deposited in the National Museum of Natural History, Smithsonian Institution, USA. Frogs from Gabon (2013) were collected during visual searches of habitat during the day and at night, as well as acoustic encounter searches at night. Frogs were handled with nitrile gloves, and individuals were collected in plastic bags. Adults were kept individually in bags, and bags were not reused. Fine-tipped rayon swabs (Dryswabs™ MW113, Medical Wire & Equipment) were used to collect samples by rubbing the ventral skin. Using a single swab, the ventral surfaces were swabbed approximately 25 times; target areas included the pelvic patch (5 passes), ventral thighs (5 passes each side with the swab) and toe webbing (5 passes on each foot). Swab samples were clipped approximately 2–3 cm from the tip and placed individually in 2 ml plastic vials. Samples were stored in 95% ethanol at ambient temperature in the field and subsequently stored at -20C in laboratory. Voucher specimens and tissue samples were deposited at the Museum of Comparative Zoology (Harvard University). Namibian frogs were detected via acoustic and visual searches conducted at night along streams within the Naukluft Mountain Zebra Park, a protected reserve in Namibia. Specimens were collected from streams originating in mountain springs on the west and east of the Naukluft Mountains. Toe tips were collected from 10 adult individuals at both the Naukluft campsite (-24.2655, 16.2395) on 2 August 2007 and at Tsams Ost (-24.2469 16.1036) on 14 August 2007 with the frogs released after sampling. Toe- clipping was first used as a sampling method for the detection of Bd until the use of skin swabbing, which is less invasive (Hyatt et al. 2007). Tissue was collected in a way to minimize potential negative effects for frogs, following the recommendations by Grafe et al. (2001). Three additional vouchers were collected in November 2007 for identification purposes and deposited in Zoologisches Museum Berlin (ZMB).

Laboratory Techniques—Various methods were used to test for the presence of Bd and are outlined below; specific information about testing, replicates, and sigmoidal amplification are outlined below. Samples were considered positive for Bd if qPCR revealed a measurable quantity (Genomic Equivalent, GE > 0.1) of Bd. The presence/absence of Bd in Namibian samples was tested by using Bd-specific primers (Bd1a/Bd2a; Annis 2004). Tissues were digested using a standard Proteinase-K protocol, and DNA was extracted using phenol-chloroform (Hillis et al. 1996). All samples were all amplified on a standard Techne 4000 PCR Thermal Cycler at the University of the Western Cape, using a positive sample of Bd culture supplied by C. Weldon. Three additional specimens were were sequenced to confirm identification (ZMB 81809 and ZMB 81812, Tsams Os; ZMB 83111, Naukluft). Sequences were submitted to GenBank as part of two previous studies (Channing and Bapista 2013; Channing et al. 2016). Positive samples were those with calculated GE over 0.1 that performed in both wells with typical sigmoid curve. DNA from samples from Cameroon were extracted using the Prepman™ Ultra Sample Preparation Reagent (Life Technologies). Real-time PCR was performed on a LightCycler 480 (Roche Applied Science, Indianapolis, IN, USA) according to the protocol of Boyle et al. (2004). A 10x dilution was made of each sample, and duplicates were used in qPCR with a standard Bd probe with Bovine serum albumin (BSA) added (Boyle et al. 2004). Standards were obtained from the Institute of Zoology, Zoological Society of London, which were made from the Bd GPL (Global Panzootic Lineage; BdGPL) lineage originally collected in Spain. Samples were considered positive if they showed distinct sigmoidal amplification in the real-time PCR, whereas negative samples and negative controls showed no such amplification (Soto-Azat et al., 2010). DNA from samples collected in Democratic Republic of the Congo (2009, 2010) and Republic of the Congo (2008) was extracted with the QIAmp DNA Mini Kit (Qiagen, Hilden, Germany) using the manufacturer’s tissue protocol. Real-time PCR was performed on a LightCycler 480 (Roche Applied Science, Indianapolis, IN, USA) according to the protocol of Boyle et al. (2004), with Bd primers specific to the ITS-1/5.8S region of the ribosomal gene. DNA extracted from a Bd-positive culture was used as a positive control and amplicons from two positive frog samples were sequenced to confirm the specificity of the amplification products. DNA from a single sample collected in Democratic Republic of the Congo in 2013 (UTEP:21111) was extracted using extracted using the Prepman™ Ultra Sample Preparation Reagent (Life Technologies), eluting in 40uL and diluting 1:10 in 0.25x TE buffer before qPCR. I ran qPCR of samples singly on an Applied Biosystems 7300 Real Time PCR system (using ITS1 and 5.8S primers) according to the methods of Boyle et. al (2004). To get the zoospore equivalents (ZE), the raw genomic equivalents output was multiplied by 80 to account for dilution during extraction. Positive controls were the Bd standards (at concentrations of 0.1, 1.0, 10, and 100 ZE) and negative controls were also used. DNA from samples collected in the Republic of the Congo (2010) was extracted from each swab using 50 μl of Prepman Ultra and detected the presence of Bd with duplicate qPCRs (Boyle et al., 2004), performed using Taqman Fast Advanced Master Mix on a ViiA7 Real-Time PCR System (Applied Biosystems, Carlsbad, CA, USA). Samples that showed signs of inhibition (non-sigmoidal amplification) were further diluted to 1:100 and re-analyzed. All samples were evaluated in duplicate plates. We generated standard curves from templates of known zoospore concentrations of Bd strain JEL427 (Puerto Rico, Luquillo) ranging from 0.1 to 1,000 zoospores (Boyle et al., 2004). To evaluate fluorescence levels of the samples and standards, we used ViiA 7 software (Applied Biosystems). DNA from samples collected in Gabon (2013) was extracted using a Qiagen DNeasy Kit (Qiagen, Hilden, Germany). Extracts were run in triplicate in qPCR assays following the fast qPCR low-volume method of Kerby et al. (2013) on a StepOne-Plus qPCR machine (Applied Biosystems). Samples were run alongside a serial dilution series of standards from 10^6 to 10^0 zoospore equivalents prepared from a live culture of BdGPL strain JEL. Samples were considered Bd positive if they amplified at or above the lowest zoospore standard (>.1 GE) in at least two of the three replicates. If only one replicate amplified, the sample was rerun in triplicate and if at least one additional replicate amplified the sample was considered Bd positive.

LITERATURE CITED Annis, S.L., F.P. Dastoor, H. Ziel, P. Daszak, and J.E. Longcore. 2004. A DNA-based assay identifies Batrachochytrium dendrobatidis in amphibians. Journal of Wildlife Diseases 40:420–428. Boyle, D.G., D.B. Boyle, V. Olsen, J.A. Morgan, and A.D. Hyatt 2004. Rapid quantitative detection of chytridiomycosis (Batrachochytrium dendrobatidis) in amphibian samples using real-time Taqman PCR assay. Diseases of aquatic organisms 60:41–148. Channing, A., and N. Baptista. 2013. Amietia angolensis and A. fuscigula (Anura: Pyxicephalidae) in southern Africa: A cold case reheated. Zootaxa 3640:501–520. Channing, A., J.H. Dehling, S. Lötters, and R. Ernst. 2016. Species boundaries and taxonomy of the African river frogs (Amphibia: Pyxicephalidae: Amietia). Zootaxa 4155:1–76. Hyatt A.D., D.G. Boyle, V. Olsen, D. B. Boyle, L. Berger, D. Obendorf, A. Dalton, К. Kriger, M. Hero, H. Hines, R. Phillott, R. Campbell, G. Marantelli, F. Gleason, and A. Colling. 2007. Diagnostic assays and sampling protocols for the detection of Batrachochytrium dendrobatidis. Diseases of aquatic organisms 73:175–192 Hillis, D.M., B.K. Mable, A. Larson, S.K. Davis, and E.A. Zimmer. 1969. Nucleic acids IV: Sequencing and cloning. In Hillis, D.M., C. Moritz, C. and B.K. Mable (eds.). Molecular Systematics 2nd edition. Sinauer Associates, USA. pp. 321–381. Kerby, J. L., A. Schieffer, J.R. Brown, and S. Whitfield. 2013. Utilization of fast qPCR techniques to detect the amphibian chytrid fungus: a cheaper and more efficient alternative method. Methods in Ecology and Evolution 4:162–166. Soto-Azat, C., B.T. Clarke, J.C. Poynton, and A.A. Cunningham. 2010. Widespread historical presence of Batrachochytridium dendrobatidis in African pipid frogs. Diversity and Distributions 16:126– 131. SUPPLEMENTAL MATERIAL S1.—List of positive African Bd records. The list shows localities that were accurate on the 30 arc sec grid. Out of the 1084 positive occurrences, a total of 490 points were used for the ENMs; 452 fell on the same grid cell as another positive record and 11 were excluded because they did not have coordinate data. The following abbreviation was used: NP = National Park. If collection date was not provided, year of publication was included and listed in italics. Records were derived from PCR or qPCR diagnostic assay using Bd primers of skin swabs or toe clips taken in field and/or excision of mouthparts from newly collected tadpoles. Records marked with an asterisk (*) indicate that Bd was detected with PCR or qPCR diagnostic assays of museum specimens. Records marked with a dagger (†) indicate that Bd was identified using morphological or histological methods, including morphological examination of tadpole mouthparts (e.g., visualization of whole- mount preparations under microscope, use of photographs of the mouth taken in the field) or histological examination of paraffin-embedded and sectioned skin tissue or toe clips that are stained to identify Bd zoosporangia with zoospores. Records marked with a double dagger (‡) indicate that Bd was not only detected, but the particular strain was identified using genotyping or genome sequencing. The Bd lineage is indicated when identified with genotyping or genome sequencing, and the total number of specimens with the lineage identified is indicated in parentheses if not all were analyzed. Hybrid/ Und. indicates that the lineage could not be identified as a hybrid or undetermined. Records without associated references that are herein reported for the first time are identified as “this study.” Taxonomy was updated to reflect that reported by Frost 2020 [46]. No. No. Country Lat Long Elevation Order Family Genus Species Year Positive Lineage Ref. 1† Botswana -24.9517 25.3429 1278 Anura Pipidae Xenopus laevis 1969 1 [42] 2† Botswana Anura Pipidae Xenopus poweri 1965 1 [42] 3‡ Burundi -3.3496 29.2731 767 Anura Arthroleptidae Leptopelis bocagii 2011 1 BdGPL [3] Hybrid/ 4‡ Burundi -3.9468 29.6274 1858 Anura Hyperoliidae Hyperolius kivuensis 2011 1 Und. [3] Hybrid/ 5‡ Burundi -3.9534 29.6147 1858 Anura Hyperoliidae Hyperolius viridiflavus 2011 2 Und. [3] Hybrid/ 6‡ Burundi -3.9468 29.6274 1858 Anura Hyperoliidae Hyperolius viridiflavus 2011 1 Und. [3] 7‡ Burundi -3.9411 29.6284 1858 Anura Pipidae Xenopus sp. 2011 1 BdGPL [3] 8‡ Burundi -3.3430 29.2731 767 Anura Ptychadenidae Ptychadena "mascareniensis" 2011 1 BdGPL [3] 9 Cameroon 2.3752 10.1088 146 Anura Arthroleptidae Arthroleptis adelphus 2014 1 [30] 10 Cameroon 4.3483 10.2326 294 Anura Arthroleptidae Arthroleptis adelphus 2014 1 [30] 11 Cameroon 2.2878 15.6706 457 Anura Arthroleptidae Arthroleptis adelphus 2014 1 [30] 12 Cameroon 2.2873 15.6726 457 Anura Arthroleptidae Arthroleptis adelphus 2014 1 [30] 13 Cameroon 6.1701 12.8272 751 Anura Arthroleptidae Arthroleptis adelphus 2014 1 [30] 14 Cameroon 4.8396 10.8696 741 Anura Arthroleptidae Arthroleptis adelphus 2015 1 [30] 15 Cameroon 2.3751 10.1087 146 Anura Arthroleptidae Arthroleptis aff. poecilonotus 2014 1 [30] 16 Cameroon 4.8368 10.8696 741 Anura Arthroleptidae Arthroleptis aff. poecilonotus 2015 1 [30] 17 Cameroon 4.8355 10.8807 773 Anura Arthroleptidae Arthroleptis aff. poecilonotus 2015 1 [30] 18 Cameroon 6.1353 9.2196 140 Anura Arthroleptidae Arthroleptis aff. poecilonotus 2015 1 [30] 2007-2008, 19 Cameroon 5.0144 9.8234 2084 Anura Arthroleptidae Arthroleptis perreti 2011 2 [8] 2007-2008, 20 Cameroon 5.0406 9.8083 1955 Anura Arthroleptidae Arthroleptis perreti 2011 1 [8] 2007-2008, 21 Cameroon 5.0406 9.8553 2046 Anura Arthroleptidae Arthroleptis perreti 2011 1 [8] 22 Cameroon 5.0406 9.8083 1843 Anura Arthroleptidae Arthroleptis perreti 2011 1 [20] 23 Cameroon 5.0406 9.8553 2046 Anura Arthroleptidae Arthroleptis perreti 2011 1 [20] 24 Cameroon 5.0144 9.8234 2084 Anura Arthroleptidae Arthroleptis perreti 2011 2 [20] BdCAPE 25‡ Cameroon 2.6310 14.0232 515 Anura Arthroleptidae Arthroleptis poecilonotus 2013 2 (1) [20], [3] 26‡ Cameroon 2.9399 11.9763 648 Anura Arthroleptidae Arthroleptis poecilonotus 2013 1 BdCAPE [20], [3] 27 Cameroon 2.9007 13.9033 576 Anura Arthroleptidae Arthroleptis poecilonotus 2013 1 [20] BdCAPE 28‡ Cameroon 2.9007 13.9033 576 Anura Arthroleptidae Arthroleptis poecilonotus 2013 12 (1) [20], [3] 29 Cameroon 4.7514 10.8243 863 Anura Arthroleptidae Arthroleptis sp. 2008 2 [8] 30 Cameroon 4.7501 10.8253 826 Anura Arthroleptidae Arthroleptis sp. 2008 3 [8] 31 Cameroon 4.7565 10.8115 816 Anura Arthroleptidae Arthroleptis sp. 2008 1 [8] 32 Cameroon 4.7514 10.8243 863 Anura Arthroleptidae Arthroleptis sp. 2008 2 [8] 33 Cameroon 6.5486 10.7600 1683 Anura Arthroleptidae Arthroleptis sp. 2011 1 [20] 34 Cameroon 4.8498 9.7718 518 Anura Arthroleptidae Arthroleptis sp. 2013 1 [20] sp. (aff. 35 Cameroon 4.1752 9.2043 1956 Anura Arthroleptidae Arthroleptis poecilonotus) 2008 1 [8] sp. (aff. 36 Cameroon 4.5570 9.5227 108 Anura Arthroleptidae Arthroleptis poecilonotus) 2008 1 [8] sp. (aff. 37 Cameroon 4.7565 10.8115 816 Anura Arthroleptidae Arthroleptis poecilonotus) 2008 1 [8] 38 Cameroon 2.2876 15.6747 457 Anura Arthroleptidae Arthroleptis sylvaticus 2014 1 [30] 39 Cameroon 2.2876 15.6813 464 Anura Arthroleptidae Arthroleptis sylvaticus 2014 1 [30] 2007-2008, 40 Cameroon 5.0382 9.7743 1466 Anura Arthroleptidae Arthroleptis tuberosus 2011 1 [8] 41 Cameroon 5.0382 9.7743 1466 Anura Arthroleptidae Arthroleptis tuberosus 2011 1 [20] 42 Cameroon 4.1752 9.2043 1956 Anura Arthroleptidae Arthroleptis variabilis 2008 1 [8] 43 Cameroon 2.9010 13.9042 576 Anura Arthroleptidae Astylosternus batesi 2013 1 [20] 44 Cameroon 4.9606 9.6686 1205 Anura Arthroleptidae Astylosternus cf. fallax 2013 2 [20] 45 Cameroon 4.9634 9.6751 1177 Anura Arthroleptidae Astylosternus cf. montanus 2013 4 [20] 46 Cameroon 4.9594 9.8651 1195 Anura Arthroleptidae Astylosternus cf. montanus 2013 1 [20] 47 Cameroon 4.9606 9.6686 1205 Anura Arthroleptidae Astylosternus cf. perreti 2013 1 [20] 48 Cameroon 4.9594 9.8651 1195 Anura Arthroleptidae Astylosternus cf. perreti 2013 8 [20] this 49* Cameroon 6.7318 11.8287 1300 Anura Arthroleptidae Astylosternus diadematus 2009 1 study this 50* Cameroon 7.7405 12.7231 1551 Anura Arthroleptidae Astylosternus diadematus 2009 1 study 51 Cameroon 4.9606 9.6686 1205 Anura Arthroleptidae Astylosternus diadematus 2013 1 [20] 52 Cameroon 6.1950 10.4616 2298 Anura Arthroleptidae Astylosternus ranoides 2008 3 [8] this 53 Cameroon 7.2522 12.0592 2004 Anura Arthroleptidae Astylosternus rheophilus 2016 17 study this 54 Cameroon 7.2575 12.0525 2118 Anura Arthroleptidae Astylosternus rheophilus 2016 1 study this 55 Cameroon 7.247 12.072 2135 Anura Arthroleptidae Astylosternus rheophilus 2016 1 study 56 Cameroon 3.1738 12.5271 632 Anura Arthroleptidae Astylosternus sp. 2013 1 [20] 57 Cameroon 4.8339 9.7783 489 Anura Arthroleptidae Cardioglossa elegans 2013 2 [20] 58* Cameroon 2.9340 11.9836 653 Anura Arthroleptidae Cardioglossa escalerae 1960 1 [20] 2007-2008, 59 Cameroon 5.0676 9.8656 1500 Anura Arthroleptidae Cardioglossa gracilis 2011 4 [8] 60 Cameroon 4.7565 10.8115 816 Anura Arthroleptidae Cardioglossa gracilis 2008 1 [8] 61 Cameroon 5.0676 9.8656 1500 Anura Arthroleptidae Cardioglossa gracilis 2011 4 [20] 62 Cameroon 5.2169 10.4857 1391 Anura Arthroleptidae Cardioglossa gracilis 2013 1 [20] 63 Cameroon 4.9594 9.8651 1195 Anura Arthroleptidae Cardioglossa gracilis 2013 1 [20] 64 Cameroon 4.3481 10.2323 294 Anura Arthroleptidae Cardioglossa leucomystax 2011 1 [8] 65 Cameroon 4.3479 10.2324 294 Anura Arthroleptidae Cardioglossa leucomystax 2014 1 [30] 2007-2008, 66 Cameroon 5.0676 9.8656 1500 Anura Arthroleptidae Cardioglossa melanogaster 2011 2 [8] 67 Cameroon 4.8593 9.8645 578 Anura Arthroleptidae Cardioglossa melanogaster 2011 1 [20] 68 Cameroon 5.0676 9.8656 1500 Anura Arthroleptidae Cardioglossa melanogaster 2011 1 [20] this 69* Cameroon 6.0928 10.3032 2146 Anura Arthroleptidae Cardioglossa oreas 2009 1 study 2007-2008, 70 Cameroon 5.0144 9.8234 2084 Anura Arthroleptidae Leptodactylodon erythrogaster 2011 1 [8] 2007-2008, 71 Cameroon 5.0307 9.8107 1946 Anura Arthroleptidae Leptodactylodon erythrogaster 2011 1 [8] 2007-2008, 72 Cameroon 5.0406 9.8553 2046 Anura Arthroleptidae Leptodactylodon erythrogaster 2011 1 [8] 73 Cameroon 5.0307 9.8107 1946 Anura Arthroleptidae Leptodactylodon erythrogaster 2011 1 [20] 74 Cameroon 5.0406 9.8553 2046 Anura Arthroleptidae Leptodactylodon erythrogaster 2011 1 [20] 75 Cameroon 5.0144 9.8234 2084 Anura Arthroleptidae Leptodactylodon erythrogaster 2011 1 [20] 2007-2008, 76 Cameroon 5.0088 9.7753 1458 Anura Arthroleptidae Leptodactylodon mertensi 2011 2 [8] 77 Cameroon 4.9594 9.8712 1194 Anura Arthroleptidae Leptodactylodon mertensi 2011 1 [20] 78 Cameroon 5.0088 9.7753 1458 Anura Arthroleptidae Leptodactylodon mertensi 2011 1 [20] 79‡ Cameroon 4.3268 9.0661 541 Anura Arthroleptidae Leptopelis aubryi 2013 1 BdCAPE [3] BdCAPE 80‡ Cameroon 4.8498 9.7718 518 Anura Arthroleptidae Leptopelis aubryi 2013 3 (1) [20], [3] 81 Cameroon 4.3223 9.0664 536 Anura Arthroleptidae Leptopelis aubryi 2013 1 [20] 82 Cameroon 4.3229 9.0827 624 Anura Arthroleptidae Leptopelis aubryi 2013 1 [20] 83 Cameroon 2.9006 13.9041 576 Anura Arthroleptidae Leptopelis aubryi 2013 2 [20] 84 Cameroon 2.3750 10.1068 119 Anura Arthroleptidae Leptopelis aubryi 2014 1 [30] 85 Cameroon 2.2850 15.6758 464 Anura Arthroleptidae Leptopelis aubryi 2014 1 [30] 86 Cameroon 4.1387 9.1983 1346 Anura Arthroleptidae Leptopelis aubryioides 2015 1 [30] 87‡ Cameroon 4.9478 9.6956 950 Anura Arthroleptidae Leptopelis boulengeri 2013 1 BdCAPE [20], [3] 88 Cameroon 4.8498 9.7718 518 Anura Arthroleptidae Leptopelis boulengeri 2013 1 [20] 89 Cameroon 4.3268 9.0661 552 Anura Arthroleptidae Leptopelis boulengeri 2013 3 [20] 2007-2008, 90 Cameroon 5.0129 9.7683 1404 Anura Arthroleptidae Leptopelis brevirostris 2011 1 [8] 91 Cameroon 5.0129 9.7683 1404 Anura Arthroleptidae Leptopelis brevirostris 2011 1 [20] 92 Cameroon 4.9634 9.6751 1177 Anura Arthroleptidae Leptopelis brevirostris 2013 1 [20] 93 Cameroon 2.3750 10.1068 119 Anura Arthroleptidae Leptopelis brevirostris 2014 1 [30] 2007-2008, 94 Cameroon 5.0243 9.7648 1417 Anura Arthroleptidae Leptopelis calcaratus 2011 2 [8] 95 Cameroon 4.9557 9.8690 1114 Anura Arthroleptidae Leptopelis calcaratus 2011 1 [20] 96 Cameroon 5.0129 9.7683 1404 Anura Arthroleptidae Leptopelis calcaratus 2011 1 [20] BdCAPE 97‡ Cameroon 4.9606 9.6686 1205 Anura Arthroleptidae Leptopelis calcaratus 2013 4 (2) [20], [3] 98 Cameroon 4.9634 9.6751 1177 Anura Arthroleptidae Leptopelis calcaratus 2013 4 [20] 99 Cameroon 4.9549 9.8672 1114 Anura Arthroleptidae Leptopelis calcaratus 2013 4 [20] 100 Cameroon 4.1390 9.1980 1346 Anura Arthroleptidae Leptopelis calcaratus 2015 2 [30] 101 Cameroon 2.6228 14.0245 513 Anura Arthroleptidae Leptopelis millsoni 2013 1 [20] 2007-2008, 102 Cameroon 5.0144 9.8234 2084 Anura Arthroleptidae Leptopelis modestus 2011 1 [8] 103 Cameroon 5.0144 9.8234 2084 Anura Arthroleptidae Leptopelis modestus 2011 1 [20] 104‡ Cameroon 5.8562 10.1589 1934 Anura Arthroleptidae Leptopelis nordequatorialis 2013 1 BdCAPE [20], [3] this 105 Cameroon 7.2635 12.0516 2018 Anura Arthroleptidae Leptopelis nordequatorialis 2016 1 study 106 Cameroon 4.7501 10.8253 826 Anura Arthroleptidae Leptopelis notatus 2008 1 [8] this 107* Cameroon 6.124 10.248 1152 Anura Arthroleptidae Leptopelis notatus 2009 1 study this 108* Cameroon 6.7318 11.8287 1300 Anura Arthroleptidae Leptopelis notatus 2009 1 study 109 Cameroon 2.6445 14.0312 524 Anura Arthroleptidae Leptopelis notatus 2013 1 [20] 110 Cameroon 2.3747 10.1079 95 Anura Arthroleptidae Leptopelis notatus 2014 1 [30] 111 Cameroon 2.4423 15.4340 516 Anura Arthroleptidae Leptopelis occelatus 2014 1 [30] 112 Cameroon 4.3505 10.2334 481 Anura Arthroleptidae Leptopelis ocellatus 2014 1 [30] 113‡ Cameroon 4.9594 9.8651 1111 Anura Arthroleptidae Leptopelis rufus 2013 1 BdCAPE [3] 114 Cameroon 4.9606 9.6686 1205 Anura Arthroleptidae Leptopelis rufus 2013 2 [20] 115 Cameroon 4.9478 9.6956 950 Anura Arthroleptidae Leptopelis rufus 2013 1 [20] BdCAPE 116‡ Cameroon 4.9594 9.8651 1195 Anura Arthroleptidae Leptopelis rufus 2013 3 (1) [20], [3] 117 Cameroon 4.8498 9.7718 518 Anura Arthroleptidae Leptopelis rufus 2013 6 [20] 118 Cameroon 4.6042 12.2045 567 Anura Arthroleptidae Leptopelis sp. 2011 1 [20] 119 Cameroon 4.9634 9.6751 1177 Anura Arthroleptidae Leptopelis sp. 2013 1 [20] 120 Cameroon 4.9478 9.6956 950 Anura Arthroleptidae Leptopelis sp. 2013 1 [20] 121 Cameroon 2.9006 13.9041 576 Anura Arthroleptidae Leptopelis sp. 2013 1 [20] 122 Cameroon 2.2823 15.6737 465 Anura Arthroleptidae Leptopelis viridis 2014 1 [30] 123 Cameroon 4.3268 9.0661 552 Anura Arthroleptidae Nyctibates corrugatus 2013 1 [20] 124 Cameroon 4.3479 10.2309 294 Anura Arthroleptidae Nyctibates corrugatus 2014 1 [30] 125 Cameroon 4.8498 9.7718 518 Anura Arthroleptidae Scotobleps gabonicus 2013 4 [20] 126 Cameroon 4.3223 9.0664 536 Anura Arthroleptidae Scotobleps gabonicus 2013 2 [20] 127 Cameroon 4.3223 9.0664 536 Anura Arthroleptidae Scotobleps gabonicus 2013 1 [20] 128 Cameroon 4.3472 10.2338 345 Anura Arthroleptidae Scotobleps gabonicus 2014 1 [30] 129 Cameroon 4.3475 10.2339 345 Anura Arthroleptidae Scotobleps gabonicus 2014 1 [30] 130 Cameroon 4.9478 9.6956 950 Anura Arthroleptidae Trichobatrachus robustus 2013 3 [20] 131 Cameroon 4.9594 9.8651 1195 Anura Arthroleptidae Trichobatrachus robustus 2013 1 [20] 132 Cameroon 4.8498 9.7718 518 Anura Arthroleptidae Trichobatrachus robustus 2013 1 [20] 133 Cameroon 2.3725 10.1042 95 Anura Bufonidae Nectophryne afra 2014 1 [30] 134‡ Cameroon 2.6106 14.0234 519 Anura Bufonidae Sclerophrys gracilipes 2013 2 BdCAPE [20], [3] 135 Cameroon 4.5570 9.5227 108 Anura Bufonidae Sclerophrys maculata 2008 1 [8] this 136* Cameroon 6.7318 11.8287 1300 Anura Bufonidae Sclerophrys maculata 2009 1 study this 137* Cameroon 7.7015 12.6767 886 Anura Bufonidae Sclerophrys maculata 2009 1 study this 138* Cameroon 7.7371 12.7212 1551 Anura Bufonidae Sclerophrys maculata 2009 1 study 139 Cameroon 4.9538 9.8661 1165 Anura Bufonidae Sclerophrys regularis 2013 2 [20] 140 Cameroon 4.3229 9.0827 624 Anura Bufonidae Sclerophrys regularis 2013 5 [20] 141 Cameroon 6.9650 10.0424 526 Anura Bufonidae Sclerophrys sp. 2011 1 [20] 142 Cameroon 6.3360 10.0351 968 Anura Bufonidae Sclerophrys sp. 2011 1 [20] 143 Cameroon 6.2307 10.0511 586 Anura Bufonidae Sclerophrys sp. 2011 1 [20] 144 Cameroon 4.6091 12.2064 568 Anura Bufonidae Sclerophrys sp. 2011 1 [20] 145 Cameroon 4.6091 12.2064 568 Anura Bufonidae Sclerophrys sp. 2011 1 [20] 146 Cameroon 3.8596 11.4956 752 Anura Bufonidae Sclerophrys sp. 2011 1 [20] 147 Cameroon 6.2020 10.4622 2230 Anura Bufonidae Wolterstorffina mirei 2008 1 [8] 148 Cameroon 4.1392 9.1110 2091 Anura Bufonidae Wolterstorffina parvipalmata 2015 1 [30] 149 Cameroon 4.1395 9.1118 2091 Anura Bufonidae Wolterstorffina parvipalmata 2015 1 [30] 150 Cameroon 4.1255 9.1459 2167 Anura Bufonidae Wolterstorffina parvipalmata 2015 1 [30] 151 Cameroon 4.1252 9.1461 2167 Anura Bufonidae Wolterstorffina parvipalmata 2015 1 [30] this 152* Cameroon 6.0488 10.4259 1213 Anura Hyperoliidae Afrixalus "quadrivittatus" 2009 1 study 153 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Afrixalus "quadrivittatus" 2011 2 [20] 154 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Afrixalus "quadrivittatus" 2011 1 [20] 155 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Afrixalus "quadrivittatus" 2011 1 [20] 156 Cameroon 5.1089 10.6159 1314 Anura Hyperoliidae Afrixalus "quadrivittatus" 2011 1 [20] 157 Cameroon 3.8596 11.4956 752 Anura Hyperoliidae Afrixalus "quadrivittatus" 2011 5 [20] 158 Cameroon 3.8596 11.4956 752 Anura Hyperoliidae Afrixalus "quadrivittatus" 2011 2 [20] 159 Cameroon 5.8562 10.1589 1934 Anura Hyperoliidae Afrixalus "quadrivittatus" 2013 1 [20] BdCAPE 160‡ Cameroon 2.9399 11.9763 648 Anura Hyperoliidae Afrixalus "quadrivittatus" 2013 5 (2) [20], [3] 161 Cameroon 2.4466 15.4328 511 Anura Hyperoliidae Afrixalus "quadrivittatus" 2014 1 [30] 162 Cameroon 4.9634 9.6751 1177 Anura Hyperoliidae Afrixalus lacteus 2013 1 [20] BdCAPE 163‡ Cameroon 4.9594 9.8651 1195 Anura Hyperoliidae Afrixalus lacteus 2013 5 (2) [20], [3] 164 Cameroon 4.9594 9.8651 1195 Anura Hyperoliidae Afrixalus laevis 2013 2 [20] 165 Cameroon 4.3223 9.0664 536 Anura Hyperoliidae Afrixalus laevis 2013 1 [20] 166 Cameroon 4.3223 9.0664 536 Anura Hyperoliidae Afrixalus laevis 2013 2 [20] 167 Cameroon 6.3360 10.0351 968 Anura Hyperoliidae Afrixalus paradorsalis 2011 1 [20] BdCAPE 168‡ Cameroon 4.8498 9.7718 518 Anura Hyperoliidae Afrixalus paradorsalis 2013 3 (1) [20], [3] 169 Cameroon 2.3751 10.1088 146 Anura Hyperoliidae Afrixalus paradorsalis 2014 1 [30] 170 Cameroon 2.2822 15.6737 465 Anura Hyperoliidae Alexteroon hypsiphonus 2014 1 [30] 171 Cameroon 2.2856 15.6748 457 Anura Hyperoliidae Alexteroon hypsiphonus 2014 1 [30] 172 Cameroon 2.2866 15.6750 464 Anura Hyperoliidae Alexteroon hypsiphonus 2014 1 [30] 173 Cameroon 2.2849 15.6753 464 Anura Hyperoliidae Alexteroon hypsiphonus 2014 1 [30] 174 Cameroon 4.3473 10.2340 345 Anura Hyperoliidae Alexteroon obstetricans 2014 1 [30] BdCAPE 175‡ Cameroon 5.8562 10.1589 1934 Anura Hyperoliidae Hyperolius ademetzi 2013 4 (3) [20], [3] 176 Cameroon 4.8498 9.7718 518 Anura Hyperoliidae Hyperolius bolifambae 2013 1 [20] 177 Cameroon 4.8498 9.7718 518 Anura Hyperoliidae Hyperolius camerunensis 2013 1 [20] 178 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Hyperolius cf. balfouri 2011 4 [20] 179 Cameroon 5.9677 10.0397 1236 Anura Hyperoliidae Hyperolius cf. balfouri 2011 3 [20] 180 Cameroon 5.8557 10.1585 1934 Anura Hyperoliidae Hyperolius cf. balfouri 2011 2 [20] 181 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Hyperolius cf. balfouri 2011 4 [20] 182 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Hyperolius cf. nasutus 2011 2 [20] 183 Cameroon 5.2169 10.4857 1391 Anura Hyperoliidae Hyperolius cf. riggenbachi 2013 1 [20] 184 Cameroon 5.2169 10.4857 1391 Anura Hyperoliidae Hyperolius cf. riggenbachi 2013 3 [20] 185 Cameroon 2.9399 11.9763 648 Anura Hyperoliidae Hyperolius cinnamomeoventris 2013 7 [20] 186 Cameroon 4.8498 9.7718 518 Anura Hyperoliidae Hyperolius concolor 2013 3 [20] 2007-2008, 187 Cameroon 5.0129 9.7683 1404 Anura Hyperoliidae Hyperolius koehleri 2011 1 [8] 188 Cameroon 4.8498 9.7718 518 Anura Hyperoliidae Hyperolius ocellatus 2013 1 [20] 189 Cameroon 2.9006 13.9041 576 Anura Hyperoliidae Hyperolius ocellatus 2013 5 [20] 190 Cameroon Anura Hyperoliidae Hyperolius riggenbachi 2008 2 [8] this 191* Cameroon 6.1162 10.2576 1207 Anura Hyperoliidae Hyperolius riggenbachi 2009 4 study this 192* Cameroon 7.7015 12.6767 886 Anura Hyperoliidae Hyperolius riggenbachi 2009 1 study this 193* Cameroon 7.7405 12.7231 1551 Anura Hyperoliidae Hyperolius riggenbachi 2009 7 study this 194* Cameroon 7.6709 12.6196 731 Anura Hyperoliidae Hyperolius riggenbachi 2009 4 study 195 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Hyperolius riggenbachi 2011 1 [20] 196 Cameroon 5.8557 10.1585 1934 Anura Hyperoliidae Hyperolius riggenbachi 2011 2 [20] 197 Cameroon 5.9090 10.1613 1543 Anura Hyperoliidae Hyperolius riggenbachi 2011 4 [20] 198 Cameroon 6.0143 10.2703 1471 Anura Hyperoliidae Hyperolius riggenbachi 2011 2 [20] 199 Cameroon 5.9677 10.0397 1236 Anura Hyperoliidae Hyperolius riggenbachi 2011 2 [20] 200 Cameroon 5.8557 10.1585 1934 Anura Hyperoliidae Hyperolius riggenbachi 2011 10 [20] 201 Cameroon 5.9090 10.1613 1543 Anura Hyperoliidae Hyperolius riggenbachi 2011 1 [20] BdCAPE 202‡ Cameroon 5.8562 10.1589 1934 Anura Hyperoliidae Hyperolius riggenbachi 2013 5 (3) [20], [3] 203 Cameroon 4.1752 9.2043 1956 Anura Hyperoliidae Hyperolius sp. 2008 2 [8] 204 Cameroon 4.8498 9.7718 518 Anura Hyperoliidae Hyperolius sp. 2013 1 [20] this 205* Cameroon 3.3913 11.4663 640 Anura Hyperoliidae Hyperolius tuberculatus 2009 1 study 206 Cameroon 5.0243 9.7648 1417 Anura Hyperoliidae Hyperolius viridistriatus 2011 1 [20] 207 Cameroon 6.5486 10.7600 1683 Anura Hyperoliidae Kassina cf. maculosa 2011 1 [20] 208 Cameroon 4.8498 9.7718 518 Anura Hyperoliidae Kassina decorata 2013 1 [20] 209‡ Cameroon 5.2509 9.5739 334 Anura Hyperoliidae Opisthothylax immaculatus 2012 1 BdCAPE [32] 210 Cameroon 2.1388 15.6557 626 Anura Hyperoliidae Phlyctimantis leonardi 2010 1 [1] 211 Cameroon 4.8339 9.7783 489 Anura Petropedetidae euskircheni 2013 2 [20] this 212* Cameroon 7.7436 12.7162 1566 Anura Petropedetidae Petropedetes sp. nov. 2009 2 study 2007-2008, 213 Cameroon Anura Phrynobatrachidae Phrynobatrachus africanus 2011 1 [8] 214 Cameroon 4.9473 9.8794 956 Anura Phrynobatrachidae Phrynobatrachus africanus 2011 1 [20] 215 Cameroon 4.8498 9.7718 518 Anura Phrynobatrachidae Phrynobatrachus africanus 2013 1 [20] 216 Cameroon 2.3743 10.1018 95 Anura Phrynobatrachidae Phrynobatrachus africanus 2014 1 [30] this 217* Cameroon 3.3913 11.4663 1566 Anura Phrynobatrachidae Phrynobatrachus auritus 2009 4 study 218 Cameroon 2.3750 10.1068 119 Anura Phrynobatrachidae Phrynobatrachus auritus 2014 2 [30] 219 Cameroon 4.3482 10.2325 294 Anura Phrynobatrachidae Phrynobatrachus auritus 2014 1 [30] 220 Cameroon 4.3472 10.2331 294 Anura Phrynobatrachidae Phrynobatrachus auritus 2014 1 [30] 221 Cameroon 4.3474 10.2338 345 Anura Phrynobatrachidae Phrynobatrachus auritus 2014 1 [30] 222 Cameroon 4.3473 10.2339 345 Anura Phrynobatrachidae Phrynobatrachus auritus 2014 1 [30] 223 Cameroon 4.8395 10.8707 741 Anura Phrynobatrachidae Phrynobatrachus auritus 2015 1 [30] 224 Cameroon 6.1352 9.2196 140 Anura Phrynobatrachidae Phrynobatrachus auritus 2015 1 [30] 225 Cameroon 6.1354 9.2199 140 Anura Phrynobatrachidae Phrynobatrachus auritus 2015 1 [30] 226 Cameroon 6.1359 9.2222 140 Anura Phrynobatrachidae Phrynobatrachus auritus 2015 1 [30] 227 Cameroon 4.7565 10.8115 816 Anura Phrynobatrachidae Phrynobatrachus calcaratus 2008 1 [8] 228 Cameroon 4.3268 9.0661 552 Anura Phrynobatrachidae Phrynobatrachus cf. auritus 2013 3 [20] 229 Cameroon 4.3268 9.0661 552 Anura Phrynobatrachidae Phrynobatrachus cornutus 2013 1 [20] 2007-2008, 230 Cameroon 5.0382 9.7743 1466 Anura Phrynobatrachidae Phrynobatrachus cricogaster 2011 1 [8] 231 Cameroon 5.0382 9.7743 1466 Anura Phrynobatrachidae Phrynobatrachus cricogaster 2011 1 [20] this 232* Cameroon 6.0488 10.4259 640 Anura Phrynobatrachidae Phrynobatrachus jimzimkusi 2009 6 study 233 Cameroon 6.1921 10.4594 2298 Anura Phrynobatrachidae Phrynobatrachus njiomock 2008 3 [8] this 234* Cameroon 6.2038 10.4596 1213 Anura Phrynobatrachidae Phrynobatrachus njiomock 2009 2 study 235 Cameroon 6.1921 10.4594 2298 Anura Phrynobatrachidae Phrynobatrachus sp. 2008 1 [8] 236 Cameroon 4.7565 10.8115 816 Anura Phrynobatrachidae Phrynobatrachus sp. 2008 3 [8] 237 Cameroon 6.2127 10.4596 2068 Anura Phrynobatrachidae Phrynobatrachus steindachneri 2008 9 [8] 238 Cameroon Anura Phrynobatrachidae Phrynobatrachus werneri 2008 4 [8] this 239 Cameroon 4.0684 9.0682 2230 Anura Pipidae Xenopus allofraseri 2016 4 study 240 Cameroon 4.3479 10.2324 294 Anura Pipidae Xenopus andrei 2014 1 [30] this 241 Cameroon 4.0681 9.0681 252 Anura Pipidae Xenopus calcaratus 2016 1 study 242* Cameroon 4.4313 14.3378 610 Anura Pipidae Xenopus cf. parafraseri 1933 1 [37] 243 Cameroon 4.1330 9.2141 975 Anura Pipidae Xenopus epitropicalis 2015 2 [30] 244 Cameroon 6.0143 10.2703 1471 Anura Pipidae Xenopus eysole 2011 1 [20] 245 Cameroon 6.5486 10.7600 1683 Anura Pipidae Xenopus eysole 2011 1 [20] 246 Cameroon 6.2405 10.5173 2086 Anura Pipidae Xenopus eysoole 2008 8 [8] 247 Cameroon Anura Pipidae Xenopus eysoole 2008 1 [8] 248 Cameroon Anura Pipidae Xenopus eysoole 2008 1 [8] this 249* Cameroon 6.0488 10.4259 252 Anura Pipidae Xenopus eysoole 2009 1 study 250 Cameroon 6.2020 10.4595 2230 Anura Pipidae Xenopus longipes 2011 7 [20] this 251 Cameroon 2.9622 12.1521 1213 Anura Pipidae Xenopus mellotropicalis 2016 3 study this 252 Cameroon 2.7795 10.8258 653 Anura Pipidae Xenopus parafraseri 2016 1 study this 253* Cameroon 7.7015 12.6767 886 Anura Pipidae Xenopus poweri 2009 4 study this 254* Cameroon 7.7436 12.7162 1566 Anura Pipidae Xenopus poweri 2009 4 study 255 Cameroon 4.5570 9.5227 108 Anura Pipidae Xenopus sp. 2008 1 [8] 256 Cameroon 6.5486 10.7600 1683 Anura Pipidae Xenopus sp. 2011 3 [20] 257 Cameroon 2.6106 14.0234 519 Anura Pipidae Xenopus sp. 2013 1 [20] this 258* Cameroon 7.7458 12.7168 1610 Anura Pipidae Xenopus sp. nov. 2009 4 study this 259 Cameroon 7.2457 12.0728 2135 Anura Pipidae Xenopus sp. nov. 2016 11 study 260 Cameroon 4.7501 10.8253 826 Anura Ptychadenidae Ptychadena "mascareniensis" 2008 1 [8] this 261* Cameroon 6.1240 10.2480 1152 Anura Ptychadenidae Ptychadena "mascareniensis" D 2009 2 study 262 Cameroon 5.5519 10.7049 1491 Anura Ptychadenidae Ptychadena sp. 2011 1 [20] 263 Cameroon 4.6116 12.2254 611 Anura Ptychadenidae Ptychadena sp. 2011 1 [20] 264 Cameroon 4.6116 12.2254 611 Anura Ptychadenidae Ptychadena sp. 2011 2 [20] 265 Cameroon 4.3268 9.0661 552 Anura Ptychadenidae Ptychadena sp. 2013 4 [20] 266 Cameroon 2.6447 14.0352 518 Anura Ranidae Amnirana albolabris 2013 1 [20] this 267* Cameroon 7.7405 12.7231 1551 Anura Ranidae Amnirana longipes 2009 1 study this 268* Cameroon 7.6709 12.6196 731 Anura Ranidae Amnirana longipes 2009 1 study 269 Cameroon 4.6042 12.2045 567 Anura Ranidae Amnirana sp. 2011 1 [20] 270 Cameroon 4.6042 12.2045 567 Anura Ranidae Amnirana sp. 2011 1 [20] BdCAPE 271‡ Cameroon 2.9006 13.9041 576 Anura Ranidae Amnirana sp. 2013 1 (1) [20], [3] 272 Cameroon 2.9010 13.9042 576 Anura Ranidae Amnirana sp. 2013 1 [20] 273 Cameroon 4.6116 12.2254 611 Anura Rhacophoridae Chiromantis rufescens 2011 1 [20] 274 Cameroon 2.6445 14.0312 524 Anura Rhacophoridae Chiromantis rufescens 2013 1 [20] 275 Cameroon 4.8498 9.7718 518 Anura Rhacophoridae Chiromantis rufescens 2013 3 [20] 276 Cameroon 4.7501 10.8253 826 Gymnophiona Dermophiidae Geotrypetes seraphini 2008 2 [8] 277 Cameroon 4.7565 10.8115 816 Gymnophiona Dermophiidae Geotrypetes seraphini 2008 6 [8] 278 Cameroon 5.3741 10.0159 732 Gymnophiona Dermophiidae Geotrypetes seraphini 2008 4 [8] 279 Cameroon 4.4144 9.4451 68 Gymnophiona Herpelidae 2008 7 [8] 280 Cameroon 4.5570 9.5227 108 Gymnophiona Herpelidae Herpele squalostoma 2008 10 [8] 281 Cameroon 4.7501 10.8253 826 Gymnophiona Herpelidae Herpele squalostoma 2008 1 [8] 282 Cameroon 4.7565 10.8115 816 Gymnophiona Indotyphlidae Idiocranium russeli 2008 18 [8] 283 Cameroon 6.22 10.46 1907 Gymnophiona Scolecomorphidae Crotaphatrema lamottei 2008 4 [8] Cameroon- this 284 Nigeria 7.0307 11.7021 1994 Anura Arthroleptidae Astylosternus rheophilus 2016 4 study Cameroon- this 285 Nigeria 7.0292 11.7058 1994 Anura Arthroleptidae Astylosternus rheophilus 2016 1 study Cameroon- this 286 Nigeria 7.0292 11.6995 1729 Anura Arthroleptidae Leptopelis nordequatorialis 2016 1 study Dem. Rep. 287 of the Congo -8.6267 27.33922 1538 Anura Arthroleptidae Arthroleptis cf. stenodactylus 2010 2 [17] Dem. Rep. 288 of the Congo -3.1455 28.9965 2084 Anura Arthroleptidae Arthroleptis sp. 2013 1 [35] Dem. Rep. 289 of the Congo -2.605 28.074 2407 Anura Arthroleptidae Arthroleptis sp. 2013 1 [35] Dem. Rep. 290 of the Congo -3.723 28.943 866 Anura Arthroleptidae Arthroleptis sp. 2014 1 [35] Dem. Rep. 291 of the Congo -5.0717 28.5958 723 Anura Arthroleptidae Arthroleptis sp. (Kabobo Plateau) 2009 1 [18] Dem. Rep. 292 of the Congo -5.164 29.113 1757 Anura Arthroleptidae Arthroleptis sp. 1 2012 1 [35] Dem. Rep. this 293 of the Congo -2.5856 16.4718 483 Anura Arthroleptidae Cardioglossa congolia 2013 1 study Dem. Rep. 294 of the Congo -3.1555 28.4211 1536 Anura Arthroleptidae Leptopelis cf. calcaratus 2010 1 [18] Dem. Rep. 295 of the Congo -2.3093 28.6494 1930 Anura Arthroleptidae Leptopelis cf. karissimbensis 2011 1 [35] Dem. Rep. this 296 of the Congo 1.3984 28.5680 777 Anura Arthroleptidae Leptopelis cf. millsoni 2009 1 study Dem. Rep. 297 of the Congo -3.3626 28.8771 2587 Anura Arthroleptidae Leptopelis karissimbensis 2008 1 [18] Dem. Rep. 298 of the Congo -3.3726 28.6779 2026 Anura Arthroleptidae Leptopelis karissimbensis 2008 1 [18] Dem. Rep. 299 of the Congo -2.2672 28.6622 2288 Anura Arthroleptidae Leptopelis karissimbensis 2008 2 [18] Dem. Rep. 300 of the Congo -3.0066 28.7501 2521 Anura Arthroleptidae Leptopelis karissimbensis 2009 1 [18] Dem. Rep. 301 of the Congo -2.5758 28.4346 1588 Anura Arthroleptidae Leptopelis karissimbensis 2009 1 [18] Dem. Rep. 302 of the Congo -3.7492 28.7474 2195 Anura Arthroleptidae Leptopelis karissimbensis 2010 1 [18] Dem. Rep. 303 of the Congo -2.3093 28.6494 1930 Anura Arthroleptidae Leptopelis karissimbensis 2011 3 [35] Dem. Rep. 304 of the Congo -2.2736 28.6630 2288 Anura Arthroleptidae Leptopelis karissimbensis 2011 3 [35] Dem. Rep. 305 of the Congo -2.2729 28.6621 2288 Anura Arthroleptidae Leptopelis kivuensis 2008 1 [18] Dem. Rep. 306 of the Congo -2.3251 28.7290 2298 Anura Arthroleptidae Leptopelis kivuensis 2011 2 [35] Dem. Rep. 307 of the Congo -2.2736 28.6630 2288 Anura Arthroleptidae Leptopelis kivuensis 2011 3 [35] Dem. Rep. 308 of the Congo -5.0649 28.9203 987 Anura Arthroleptidae Leptopelis kivuensis 2012 1 [35] Dem. Rep. 309 of the Congo -3.520 28.419 1156 Anura Arthroleptidae Leptopelis kivuensis 2013 1 [35] Dem. Rep. 310 of the Congo -2.589 28.051 1238 Anura Arthroleptidae Leptopelis kivuensis 2013 1 [35] Dem. Rep. 311 of the Congo -3.3726 28.6432 1800 Anura Arthroleptidae Leptopelis mtoewaate 2008 1 [18] Dem. Rep. 312 of the Congo -3.1555 28.4211 1536 Anura Arthroleptidae Leptopelis mtoewaate 2010 2 [18] Dem. Rep. 313 of the Congo -3.1572 28.4234 1536 Anura Arthroleptidae Leptopelis mtoewaate 2010 1 [18] Dem. Rep. 314 of the Congo -8.6267 27.3392 1538 Anura Arthroleptidae Leptopelis parbocagii 2010 1 [17] Dem. Rep. 315 of the Congo -2.2736 28.6630 2288 Anura Arthroleptidae Leptopelis sp. 2011 2 [35] Dem. Rep. 316 of the Congo -3.520 28.419 1156 Anura Arthroleptidae Leptopelis sp. 2013 2 [35] Dem. Rep. 317 of the Congo -3.1539 28.9937 2045 Anura Arthroleptidae Leptopelis sp. 2013 2 [35] Dem. Rep. 318 of the Congo -1.925 28.019 1075 Anura Arthroleptidae Leptopelis sp. 2013 1 [35] Dem. Rep. 319 of the Congo -3.220 28.656 1891 Anura Arthroleptidae Leptopelis sp. 2014 1 [35] Dem. Rep. 320 of the Congo -3.383 28.567 1460 Anura Arthroleptidae Leptopelis sp. 2014 1 [35] Dem. Rep. sp. 1 (Mt. 321 of the Congo -0.1261 29.4328 2807 Anura Arthroleptidae Leptopelis Tshiaberimu) 2008 1 [18] Dem. Rep. 322 of the Congo -3.0288 28.2824 1144 Anura Bufonidae Nectophryne cf. batesii 2010 1 [18] Dem. Rep. 323 of the Congo -3.44 28.351 1142 Anura Bufonidae Sclerophrys sp. 2013 1 [35] Dem. Rep. 324 of the Congo -3.8756 27.9252 797 Anura Hyperoliidae Afrixalus "quadrivittatus" 2012 1 [35] Dem. Rep. 325 of the Congo -3.4039 28.5236 1413 Anura Hyperoliidae Afrixalus cf. "quadrivittatus" 1 2008 1 [18] Dem. Rep. 326 of the Congo 0.6649 29.8781 852 Anura Hyperoliidae Afrixalus cf. "quadrivittatus" 2 2008 1 [18] Dem. Rep. 327 of the Congo -2.2289 28.7802 1987 Anura Hyperoliidae Afrixalus cf. laevis 2009 1 [18] Dem. Rep. 328 of the Congo -2.3251 28.7290 2298 Anura Hyperoliidae Afrixalus cf. laevis 2011 4 [35] Dem. Rep. 329 of the Congo -2.2736 28.6630 2288 Anura Hyperoliidae Afrixalus cf. laevis 2011 1 [35] Dem. Rep. 330 of the Congo -1.674 28.128 687 Anura Hyperoliidae Afrixalus osorioi 2013 2 [35] Dem. Rep. 331 of the Congo -1.782 27.533 1081 Anura Hyperoliidae Afrixalus osorioi 2013 1 [35] Dem. Rep. 332 of the Congo -2.558 27.994 723 Anura Hyperoliidae Afrixalus osorioi 2013 1 [35] Dem. Rep. 333 of the Congo -3.519 28.419 1156 Anura Hyperoliidae Afrixalus sp. 2013 2 [35] Dem. Rep. 334 of the Congo -3.445 28.356 1174 Anura Hyperoliidae Afrixalus sp. 2013 1 [35] Dem. Rep. 335 of the Congo -2.558 27.994 723 Anura Hyperoliidae Afrixalus sp. 2013 2 [35] Dem. Rep. 336 of the Congo -1.637 27.115 590 Anura Hyperoliidae Afrixalus sp. 2013 1 [35] Dem. Rep. 337 of the Congo -1.384 27.198 768 Anura Hyperoliidae Afrixalus sp. 2013 1 [35] Dem. Rep. 338 of the Congo -3.1447 28.9986 2084 Anura Hyperoliidae Afrixalus sp. 1 2012 1 [35] Dem. Rep. 339 of the Congo -3.1447 28.9986 2084 Anura Hyperoliidae Afrixalus sp. 3 2012 1 [35] Dem. Rep. 340 of the Congo -3.44 28.351 1142 Anura Hyperoliidae Afrixalus wittei 2013 2 [35] Dem. Rep. 341 of the Congo -1.637 27.115 590 Anura Hyperoliidae Afrixalus wittei 2013 1 [35] Dem. Rep. 342 of the Congo -3.3726 28.6779 2026 Anura Hyperoliidae Callixalus pictus 2008 1 [18] Dem. Rep. 343 of the Congo -3.0885 28.8104 2786 Anura Hyperoliidae Chrysobatrachus cupreonitens 2009 1 [18] Dem. Rep. 344 of the Congo -3.0766 28.8167 2915 Anura Hyperoliidae Chrysobatrachus cupreonitens 2009 1 [18] Dem. Rep. 345 of the Congo -3.4318 29.0141 2868 Anura Hyperoliidae Chrysobatrachus cupreonitens 2011 1 [18] Dem. Rep. 346 of the Congo -3.4315 29.0116 2868 Anura Hyperoliidae Chrysobatrachus cupreonitens 2011 1 [18] Dem. Rep. this 347 of the Congo 1.3969 28.5859 760 Anura Hyperoliidae Hyperolius bolifambae 2009 1 study Dem. Rep. 348 of the Congo -0.1255 29.4335 2796 Anura Hyperoliidae Hyperolius castaneus 2008 1 [18] Dem. Rep. 349 of the Congo -2.3251 28.7290 2298 Anura Hyperoliidae Hyperolius castaneus 2011 5 [35] Dem. Rep. 350 of the Congo -2.2736 28.6630 2288 Anura Hyperoliidae Hyperolius castaneus 2011 2 [35] Dem. Rep. 351 of the Congo -3.1529 28.9920 2045 Anura Hyperoliidae Hyperolius castaneus 2013 1 [35] Dem. Rep. 352 of the Congo -3.1486 28.9934 2084 Anura Hyperoliidae Hyperolius castaneus 2013 1 [35] Dem. Rep. 353 of the Congo -3.1459 29.0005 2051 Anura Hyperoliidae Hyperolius castaneus 2013 4 [35] Dem. Rep. 354 of the Congo -3.1454 28.9990 2084 Anura Hyperoliidae Hyperolius castaneus 2013 1 [35] Dem. Rep. cf. this 355 of the Congo -3.3403 28.1309 834 Anura Hyperoliidae Hyperolius cinnamomeoventris 2010 1 study Dem. Rep. 356 of the Congo -8.7419 28.2167 1009 Anura Hyperoliidae Hyperolius cf. kivuensis 2010 1 [17] Dem. Rep. 357 of the Congo -3.0176 28.3342 1233 Anura Hyperoliidae Hyperolius cf. langi 2008 1 [18] Dem. Rep. 358 of the Congo -3.3734 28.6431 1800 Anura Hyperoliidae Hyperolius cf. tuberculatus 2008 1 [18] Dem. Rep. 359 of the Congo -3.4039 28.5866 1908 Anura Hyperoliidae Hyperolius cf. tuberculatus 2008 1 [18] Dem. Rep. 360 of the Congo -3.0066 28.7501 2521 Anura Hyperoliidae Hyperolius constellatus 2009 1 [18] Dem. Rep. 361 of the Congo -3.0766 28.8167 2915 Anura Hyperoliidae Hyperolius constellatus 2009 2 [18] Dem. Rep. 362 of the Congo -5.1077 29.0413 2402 Anura Hyperoliidae Hyperolius constellatus 2009 1 [18] Dem. Rep. 363 of the Congo -3.1486 28.9934 2084 Anura Hyperoliidae Hyperolius constellatus 2013 1 [35] Dem. Rep. 364 of the Congo -2.3251 28.7290 2298 Anura Hyperoliidae Hyperolius discodactylus 2011 1 [35] Dem. Rep. 365 of the Congo -3.8750 28.0303 1289 Anura Hyperoliidae Hyperolius frontalis 2012 1 [35] Dem. Rep. 366 of the Congo -3.0386 28.4334 1298 Anura Hyperoliidae Hyperolius hutsebauti 2008 1 [18] Dem. Rep. 367† of the Congo -2.3128 28.7552 2161 Anura Hyperoliidae Hyperolius kivuensis 2007 1 [16] Dem. Rep. 368 of the Congo 0.6649 29.8781 852 Anura Hyperoliidae Hyperolius kivuensis 2008 1 [18] Dem. Rep. 369 of the Congo -2.6859 28.8683 1569 Anura Hyperoliidae Hyperolius kivuensis 2009 1 [18] Dem. Rep. 370 of the Congo -4.2841 28.9633 986 Anura Hyperoliidae Hyperolius kivuensis 2009 1 [18] Dem. Rep. 371 of the Congo -2.2275 28.7810 1987 Anura Hyperoliidae Hyperolius kivuensis 2009 1 [18] Dem. Rep. 372† of the Congo -1.8873 28.4495 976 Anura Hyperoliidae Hyperolius langi 2007 1 [16] Dem. Rep. this 373 of the Congo 1.4092 28.5723 768 Anura Hyperoliidae Hyperolius langi 2009 1 study Dem. Rep. 374 of the Congo -7.7149 29.7696 2014 Anura Hyperoliidae Hyperolius nasicus 2010 2 [17] Dem. Rep. 375 of the Congo -3.1352 28.9970 2010 Anura Hyperoliidae Hyperolius nasutus 2013 1 [35] Dem. Rep. 376 of the Congo -7.7094 29.7810 2010 Anura Hyperoliidae Hyperolius quinquevittatus 2010 1 [17] Dem. Rep. 377 of the Congo -8.4058 27.2821 1359 Anura Hyperoliidae Hyperolius quinquevittatus 2010 1 [17] Dem. Rep. 378 of the Congo -3.440 28.351 1142 Anura Hyperoliidae Hyperolius robustus 2013 1 [35] Dem. Rep. 379 of the Congo -3.296 28.331 1300 Anura Hyperoliidae Hyperolius sp. 2013 3 [35] Dem. Rep. 380 of the Congo -3.519 28.419 1156 Anura Hyperoliidae Hyperolius sp. 2013 2 [35] Dem. Rep. 381 of the Congo -3.1529 28.9920 2045 Anura Hyperoliidae Hyperolius sp. 2013 1 [35] Dem. Rep. 382 of the Congo -3.1454 28.9990 2084 Anura Hyperoliidae Hyperolius sp. 2013 2 [35] Dem. Rep. 383 of the Congo -3.1377 28.9919 2010 Anura Hyperoliidae Hyperolius sp. 2013 2 [35] Dem. Rep. 384 of the Congo -3.1352 28.9970 2010 Anura Hyperoliidae Hyperolius sp. 2013 3 [35] Dem. Rep. 385 of the Congo -3.445 28.356 1174 Anura Hyperoliidae Hyperolius sp. 2013 1 [35] Dem. Rep. 386 of the Congo -3.44 28.351 1142 Anura Hyperoliidae Hyperolius sp. 2013 5 [35] Dem. Rep. 387 of the Congo -1.757 27.449 1152 Anura Hyperoliidae Hyperolius sp. 2013 1 [35] Dem. Rep. 388 of the Congo -2.558 27.994 723 Anura Hyperoliidae Hyperolius sp. 2013 1 [35] Dem. Rep. 389 of the Congo -5.24 29.105 2003 Anura Hyperoliidae Hyperolius sp. 1 2012 1 [35] Dem. Rep. 390 of the Congo -5.152 28.744 884 Anura Hyperoliidae Hyperolius sp. 1 2012 1 [35] Dem. Rep. 391 of the Congo -3.1447 28.9986 2084 Anura Hyperoliidae Hyperolius sp. 2 2012 1 [35] Dem. Rep. 392 of the Congo -3.8557 28.0411 1427 Anura Hyperoliidae Hyperolius sp. 4 2012 1 [35] Dem. Rep. 393 of the Congo -3.1980 28.9924 2122 Anura Hyperoliidae Hyperolius sp. 5 2012 1 [35] Dem. Rep. 394 of the Congo -3.1980 28.9924 2122 Anura Hyperoliidae Hyperolius sp. 7 2012 1 [35] Dem. Rep. 395 of the Congo -7.7149 29.7696 2014 Anura Hyperoliidae Kassina cf. senegalensis 2010 2 [17] Dem. Rep. 396 of the Congo -3.3733 28.6432 1800 Anura Hyperoliidae Phlyctimantis verrucosus 2008 2 [18] Dem. Rep. 397 of the Congo -3.9018 28.0192 1205 Anura Hyperoliidae Phlyctimantis verrucosus 2012 1 [35] Dem. Rep. 398 of the Congo -2.2634 28.6678 2390 Anura Phrynobatrachidae Phrynobatrachus acutirostris 2008 1 [18] Dem. Rep. 399* of the Congo Anura Phrynobatrachidae Phrynobatrachus asper 1950 1 [35] Dem. Rep. 400 of the Congo -3.0781 28.7679 3117 Anura Phrynobatrachidae Phrynobatrachus asper 2009 1 [18] Dem. Rep. 401 of the Congo -0.1238 29.4317 2639 Anura Phrynobatrachidae Phrynobatrachus cf. graueri 2008 1 [18] Dem. Rep. 402 of the Congo -5.0600 28.9215 987 Anura Phrynobatrachidae Phrynobatrachus cf. natalensis 1 2009 1 [18] Dem. Rep. 403 of the Congo -3.0368 28.4223 1351 Anura Phrynobatrachidae Phrynobatrachus cf. natalensis 1 2010 1 [18] Dem. Rep. 404 of the Congo -8.7190 27.4227 1440 Anura Phrynobatrachidae Phrynobatrachus cf. natalensis 2 2010 1 [17] Dem. Rep. 405 of the Congo -3.0288 28.2824 1144 Anura Phrynobatrachidae Phrynobatrachus dendrobates 2010 2 [18] Dem. Rep. 406 of the Congo -3.1578 28.4341 1507 Anura Phrynobatrachidae Phrynobatrachus dendrobates 2010 1 [18] Dem. Rep. 407 of the Congo -2.350 28.081 1362 Anura Phrynobatrachidae Phrynobatrachus krefftii 2013 1 [35] Dem. Rep. 408 of the Congo -3.0176 28.3342 1233 Anura Phrynobatrachidae Phrynobatrachus sp. 2008 1 [18] Dem. Rep. 409 of the Congo -5.0600 28.9215 987 Anura Phrynobatrachidae Phrynobatrachus sp. 2009 1 [18] Dem. Rep. 410 of the Congo -4.6799 28.6534 790 Anura Phrynobatrachidae Phrynobatrachus sp. 2009 1 [18] Dem. Rep. 411 of the Congo -1.844 27.608 1114 Anura Phrynobatrachidae Phrynobatrachus sp. 2013 1 [35] Dem. Rep. 412 of the Congo -3.234 28.519 1942 Anura Phrynobatrachidae Phrynobatrachus sp. 2014 1 [35] Dem. Rep. 413 of the Congo -3.253 28.543 1910 Anura Phrynobatrachidae Phrynobatrachus sp. 2014 1 [35] Dem. Rep. 414 of the Congo -3.818 28.969 2224 Anura Phrynobatrachidae Phrynobatrachus sp. 2014 5 [35] Dem. Rep. 415 of the Congo -5.258 28.888 1290 Anura Phrynobatrachidae Phrynobatrachus sp. 1 2012 1 [35] Dem. Rep. 416 of the Congo -5.080 28.734 1017 Anura Phrynobatrachidae Phrynobatrachus sp. 1 2012 2 [35] Dem. Rep. 417 of the Congo -2.2675 28.6621 2288 Anura Phrynobatrachidae Phrynobatrachus versicolor 2008 1 [18] Dem. Rep. 418 of the Congo -7.7094 29.7810 2010 Anura Pipidae Xenopus laevis 2010 1 [17] Dem. Rep. 419 of the Congo -2.3093 28.6494 1930 Anura Pipidae Xenopus sp. 2011 1 [35] Dem. Rep. 420 of the Congo -3.1529 28.9920 2045 Anura Pipidae Xenopus sp. 2013 1 [35] Dem. Rep. 421 of the Congo -2.3093 28.6494 1930 Anura Pipidae Xenopus wittei 2011 2 [35] Dem. Rep. 422 of the Congo -8.4741 27.3154 1581 Anura Ptychadenidae Ptychadena cf. uzungwensis 2010 1 [17] Dem. Rep. 423 of the Congo -3.8756 27.9252 797 Anura Ptychadenidae Ptychadena chrysogaster 2012 1 [35] Dem. Rep. 424 of the Congo -3.8756 27.9252 797 Anura Ptychadenidae Ptychadena sp. 1 2012 1 [35] Dem. Rep. 425 of the Congo -5.152 28.744 884 Anura Ptychadenidae Ptychadena sp. 1 2012 1 [35] Dem. Rep. 426 of the Congo -5.149 28.742 874 Anura Ptychadenidae Ptychadena sp. 1 2012 1 [35] Dem. Rep. 427 of the Congo -5.0649 28.9157 975 Anura Ptychadenidae Ptychadena sp. A 2012 1 [35] Dem. Rep. 428 of the Congo -2.2736 28.6630 2288 Anura Pyxicephalidae Amietia sp. 2011 1 [35] Dem. Rep. 429 of the Congo -3.1459 28.9950 2084 Anura Pyxicephalidae Amietia sp. 2013 1 [35] Dem. Rep. 430 of the Congo -7.6188 29.7866 2032 Anura Pyxicephalidae Amietia sp. 1 2010 1 [17] Dem. Rep. 431 of the Congo -7.7094 29.7810 2010 Anura Pyxicephalidae Amietia sp. 1 2010 1 [17] Dem. Rep. 432 of the Congo -5.0569 28.9236 960 Anura Pyxicephalidae Amietia sp. 1 (Force Bendera) 2009 2 [18] Dem. Rep. 433 of the Congo -3.3726 28.6779 2026 Anura Pyxicephalidae Amietia sp. 2 (Itombwe) 2008 1 [18] Dem. Rep. 434 of the Congo -3.0766 28.8167 2915 Anura Pyxicephalidae Amietia sp. 2 (Itombwe) 2009 1 [18] Dem. Rep. 435 of the Congo -2.9969 28.8884 2787 Anura Pyxicephalidae Amietia sp. 2 (Itombwe) 2009 2 [18] Dem. Rep. 436 of the Congo -2.9969 28.8884 2787 Anura Pyxicephalidae Amietia sp. 2 (Itombwe) 2009 1 [18] Dem. Rep. 437 of the Congo -3.4314 29.0115 2868 Anura Pyxicephalidae Amietia sp. 2 (Itombwe) 2011 1 [18] Dem. Rep. 438 of the Congo -8.4658 27.3307 1468 Anura Pyxicephalidae Amietia sp. 3 2010 1 [17] Equatorial 439* Guinea 3.628 8.803 1805 Anura Arthroleptidae Arthroleptis bioko 1998 1 [23] Equatorial 440* Guinea 3.696 8.871 649 Anura Arthroleptidae Arthroleptis poecilonotus 1998 1 [23] Equatorial 441* Guinea 3.325 8.671 1175 Anura Arthroleptidae Leptopelis calcaratus 1998 6 [23] Equatorial 442* Guinea Anura Bufonidae Sclerophrys camerunensis 1966 1 [23] Equatorial 443* Guinea 3.361 8.662 1450 Anura Bufonidae Sclerophrys camerunensis 1998 4 [23] Equatorial 444* Guinea 3.325 8.671 1175 Anura Hyperoliidae Afrixalus laevis 1998 6 [23] Equatorial 445* Guinea 3.461 8.552 -9999 Anura Hyperoliidae Afrixalus paradorsalis 1998 1 [23] Equatorial 446* Guinea 3.355 8.622 1826 Anura Hyperoliidae Hyperolius endjami 1998 2 [23] Equatorial BdGPL 447‡ Guinea 3.325 8.671 1175 Anura Hyperoliidae Hyperolius endjami 1998 1 (1) [23], [3] Equatorial 448* Guinea 3.355 8.622 1826 Anura Hyperoliidae Hyperolius tuberculatus 1998 3 [23] Equatorial 449* Guinea 3.367 8.665 1503 Anura Phrynobatrachidae Phrynobatrachus auritus 1998 1 [23] Equatorial 450* Guinea 3.331 8.667 1175 Anura Phrynobatrachidae Phrynobatrachus auritus 1998 3 [23] Equatorial 451* Guinea 3.332 8.67 1175 Anura Phrynobatrachidae Phrynobatrachus auritus 1998 1 [23] Equatorial 452* Guinea 3.483 8.582 -9999 Anura Phrynobatrachidae Phrynobatrachus cornutus 1998 2 [23] Equatorial 453* Guinea 3.325 8.671 1175 Anura Phrynobatrachidae Phrynobatrachus cornutus 1998 1 [23] Equatorial 454* Guinea 3.325 8.671 1175 Anura Phrynobatrachidae Phrynobatrachus cornutus 1998 1 [23] Equatorial 455* Guinea 3.331 8.667 1175 Anura Phrynobatrachidae Phrynobatrachus cornutus 1998 10 [23] Equatorial 456* Guinea 3.696 8.871 649 Anura Phrynobatrachidae Phrynobatrachus cornutus 1998 1 [23] Equatorial 457* Guinea 3.52 8.586 46 Anura Pipidae Xenopus allofraseri 1998 2 [23] Equatorial 458* Guinea 3.52 8.586 46 Anura Pipidae Xenopus calcaratus 1998 2 [23] Equatorial 459* Guinea 3.528 8.579 20 Anura Pipidae Xenopus calcaratus 1998 1 [23] Equatorial 460* Guinea 3.711 8.667 55 Anura Ranidae Amnirana albolabris 1998 1 [23] Equatorial 461* Guinea 3.461 8.552 -9999 Anura Ranidae Amnirana albolabris 1998 4 [23] Equatorial 462* Guinea 3.468 8.493 -9999 Anura Ranidae Amnirana albolabris 1998 1 [23] 463† Eswatini Anura Pipidae Xenopus muelleri 1988 1 [42] 464† Eswatini Anura Pipidae Xenopus muelleri 1991 1 [42] 465 Ethiopia 6.7170 39.7257 2452 Anura Arthroleptidae Leptopelis cf. argenteus 2009 4 [14] 466 Ethiopia 6.6386 39.7339 1918 Anura Arthroleptidae Leptopelis cf. ragazzii 2009 1 [14] 467 Ethiopia 6.7713 39.7245 2910 Anura Arthroleptidae Leptopelis gramineus 2008 2 [14] 468 Ethiopia 6.7591 39.7214 2818 Anura Arthroleptidae Leptopelis gramineus 2008 2 [14] 469 Ethiopia 6.7164 39.7257 2385 Anura Arthroleptidae Leptopelis gramineus 2008 2 [14] 470 Ethiopia 6.7713 39.7245 2910 Anura Arthroleptidae Leptopelis ragazzii 2008 6 [14] 471 Ethiopia 6.7170 39.7257 2452 Anura Arthroleptidae Leptopelis ragazzii 2009 2 [14] 472 Ethiopia 6.7713 39.7245 2910 Anura Arthroleptidae Leptopelis ragazzii 2009 3 [14] 473‡ Ethiopia 6.7572 39.7212 2824 Anura Arthroleptidae Leptopelis sp. 2011 1 BdGPL [32] 474 Ethiopia 7.1702 36.2717 1564 Anura Arthroleptidae Leptopelis vannutellii 2009 2 [14] 475 Ethiopia 7.5019 36.0918 2190 Anura Arthroleptidae Leptopelis vannutellii 2009 2 [14] 476 Ethiopia 6.7576 39.7500 3248 Anura Brevicipitidae Balebreviceps hillmani 2009 1 [14] 477 Ethiopia 6.7776 39.7455 3272 Anura Brevicipitidae Balebreviceps hillmani 2009 1 [14] 478 Ethiopia 6.7735 39.7278 3014 Anura Bufonidae Altiphrynoides malcolmi 2008 1 [14] 479 Ethiopia 6.7576 39.7500 3248 Anura Bufonidae Altiphrynoides malcolmi 2009 1 [14] 480 Ethiopia 7.2493 36.2554 1709 Anura Hyperoliidae Afrixalus clarkei 2009 3 [14] 481 Ethiopia 7.3180 36.0782 1960 Anura Hyperoliidae Afrixalus enseticola 2009 1 [14] 482‡ Ethiopia 6.6865 39.6641 1866 Anura Hyperoliidae Afrixalus enseticola 2011 1 BdGPL [32] 483 Ethiopia 6.7170 39.7257 2452 Anura Hyperoliidae Afrixalus sp. 2009 1 [14] 484 Ethiopia 7.2493 36.2554 1709 Anura Hyperoliidae Hyperolius cf. kivuensis 2009 2 [14] 485 Ethiopia 7.3021 36.1164 1975 Anura Hyperoliidae Hyperolius viridiflavus 2009 1 [14] 486 Ethiopia 7.2493 36.2554 1709 Anura Hyperoliidae Paracassina obscura 2009 1 [14] 487 Ethiopia 6.7576 39.7500 3248 Anura Petropedetidae Ericabatrachus baleensis 2009 2 [14] 488 Ethiopia 7.3101 36.1205 1893 Anura Phrynobatrachidae Phrynobatrachus cf. minutus 2009 1 [14] 489 Ethiopia 7.2493 36.2554 1709 Anura Phrynobatrachidae Phrynobatrachus minutus 2009 2 [14] 490 Ethiopia 7.3101 36.1205 1893 Anura Phrynobatrachidae Phrynobatrachus minutus 2009 3 [14] 491 Ethiopia 7.2672 36.2590 1747 Anura Phrynobatrachidae Phrynobatrachus natalensis 2009 1 [14] 492 Ethiopia 6.6386 39.7339 1918 Anura Pipidae Xenopus clivii 2009 1 [14] 493 Ethiopia 6.7733 39.7254 3014 Anura Ptychadenidae Ptychadena erlangeri 2008 1 [14] 494 Ethiopia 6.7676 39.7222 2910 Anura Ptychadenidae Ptychadena erlangeri 2008 1 [14] 495 Ethiopia 6.7530 39.7190 2798 Anura Ptychadenidae Ptychadena erlangeri 2008 1 [14] 496 Ethiopia 6.7713 39.7245 2910 Anura Ptychadenidae Ptychadena neumanni 2008 1 [14] 497 Ethiopia 6.7596 39.7215 2818 Anura Ptychadenidae Ptychadena neumanni 2009 2 [14] 498 Ethiopia 7.2672 36.2590 1747 Anura Ptychadenidae Ptychadena neumanni 2009 1 [14] 499 Ethiopia 7.272 36.26 1747 Anura Ptychadenidae Ptychadena neumanni 2009 2 [14] 500 Ethiopia 7.31005 36.1205 1893 Anura Ptychadenidae Ptychadena neumanni 2009 1 [14] this 501 Gabon -1.6278 12.3631 752 Anura Arthroleptidae Arthroleptis adelphus 2013 1 study this 502 Gabon 1.1544 12.6992 505 Anura Arthroleptidae Arthroleptis adelphus 2013 1 study this 503 Gabon -1.6278 12.3631 752 Anura Arthroleptidae Arthroleptis cf. poecilonotus 2013 1 study this 504 Gabon -0.3636 11.7872 259 Anura Arthroleptidae Arthroleptis cf. poecilonotus 2013 1 study 505 Gabon 0.5112 12.8028 506 Anura Arthroleptidae Arthroleptis poecilonotus 2009 1 [2] 506 Gabon -0.8683 12.6724 278 Anura Arthroleptidae Arthroleptis poecilonotus 2015 2 [26] 507 Gabon 0.511 12.803 506 Anura Arthroleptidae Arthroleptis sp. 2010 1 [23] 508 Gabon 0.4536 10.2781 136 Anura Arthroleptidae Arthroleptis sp. A 2009 1 [2] 509 Gabon -0.1684 10.7829 36 Anura Arthroleptidae Arthroleptis sylvaticus 2015 1 [26] this 510 Gabon -0.3894 11.8239 243 Anura Arthroleptidae Arthroleptis sylvaticus group 2013 1 study this 511 Gabon 1.1544 12.6992 505 Anura Arthroleptidae Arthroleptis sylvaticus group 2013 2 study this 512 Gabon -1.8553 13.8542 407 Anura Arthroleptidae Arthroleptis sylvaticus group 2013 1 study this 513 Gabon -1.8553 13.8542 407 Anura Arthroleptidae Arthroleptis sylvaticus group 2013 1 study this 514 Gabon -1.8553 13.8542 407 Anura Arthroleptidae Arthroleptis sylvaticus group 2013 1 study this 515 Gabon 1.1544 12.6992 505 Anura Arthroleptidae Arthroleptis variabilis 2013 1 study 516 Gabon -0.1766 10.7797 54 Anura Arthroleptidae Arthroleptis variabilis 2015 1 [26] 517 Gabon -0.0521 11.1649 194 Anura Arthroleptidae Arthroleptis variabilis 2015 1 [26] this 518 Gabon -1.6278 12.3631 752 Anura Arthroleptidae Astylosternus batesi 2013 2 study this 519 Gabon -1.8553 13.8542 407 Anura Arthroleptidae Astylosternus batesi 2013 1 study 520 Gabon 0.4536 10.2781 136 Anura Arthroleptidae Cardioglossa elegans 2009 1 [2] this 521 Gabon -1.6278 12.3631 752 Anura Arthroleptidae Cardioglossa gracilis 2013 1 study this 522 Gabon 1.1544 12.6992 505 Anura Arthroleptidae Cardioglossa leucomystax 2013 1 study 523 Gabon 0.5112 12.8028 506 Anura Arthroleptidae Leptopelis aubryi 2009 1 [2] 524 Gabon -0.0554 11.1664 194 Anura Arthroleptidae Leptopelis aubryi 2015 1 [26] 525 Gabon 0.506 12.796 532 Anura Arthroleptidae Leptopelis boulengeri 2010 1 [23] this 526 Gabon 1.1544 12.6992 505 Anura Arthroleptidae Leptopelis boulengeri 2013 1 study 527 Gabon 0.4496 10.2758 130 Anura Arthroleptidae Leptopelis brevirostris 2009 1 [2] 528 Gabon 0.6256 10.3965 494 Anura Arthroleptidae Leptopelis brevirostris 2009 1 [2] 529 Gabon 0.4999 12.8018 493 Anura Arthroleptidae Leptopelis calcaratus 2009 1 [2] 530 Gabon 0.5112 12.8028 506 Anura Arthroleptidae Leptopelis calcaratus 2009 1 [2] 531 Gabon -0.1809 10.7783 54 Anura Arthroleptidae Leptopelis millsoni 2015 1 [26] 532 Gabon 0.5112 12.8028 506 Anura Arthroleptidae Leptopelis notatus 2009 1 [2] 533 Gabon 0.506 12.796 532 Anura Arthroleptidae Leptopelis notatus 2010 1 [23] this 534 Gabon -1.8553 13.8542 407 Anura Arthroleptidae Leptopelis notatus 2013 1 study 535 Gabon 0.4999 12.8018 493 Anura Arthroleptidae Leptopelis ocellatus 2009 1 [2] 536 Gabon 0.5037 12.7941 532 Anura Arthroleptidae Leptopelis ocellatus 2009 2 [2] 537 Gabon -0.226 12.298 213 Anura Arthroleptidae Leptopelis ocellatus 2011 1 [23] 538 Gabon -0.189 12.185 213 Anura Arthroleptidae Leptopelis ocellatus 2011 3 [23] this 539 Gabon -1.6278 12.3631 752 Anura Arthroleptidae Leptopelis ocellatus 2013 1 study this 540 Gabon -0.3636 11.7872 259 Anura Arthroleptidae Leptopelis ocellatus 2013 1 study 541 Gabon -0.8683 12.6724 278 Anura Arthroleptidae Leptopelis ocellatus 2015 1 [26] 542 Gabon -0.8683 12.6724 278 Anura Arthroleptidae Leptopelis ocellatus 2015 1 [26] 543 Gabon 0.6233 10.4028 496 Anura Arthroleptidae Leptopelis rufus 2009 1 [2] 544 Gabon -0.0622 11.1587 102 Anura Arthroleptidae Leptopelis rufus 2015 1 [26] 545 Gabon 0.4536 10.2781 136 Anura Arthroleptidae Scotobleps gabonicus 2009 1 [2] 546 Gabon 0.6212 10.40763 496 Anura Arthroleptidae Scotobleps gabonicus 2009 1 [2] 547 Gabon 0.6256 10.3965 494 Anura Arthroleptidae Scotobleps gabonicus 2009 1 [2] this 548 Gabon -0.3636 11.7872 259 Anura Arthroleptidae Scotobleps gabonicus 2013 2 study 549 Gabon -0.0492 11.1643 174 Anura Arthroleptidae Scotobleps gabonicus 2015 1 [26] 550 Gabon -0.6407 10.2187 40 Anura Arthroleptidae Scotobleps gabonicus 2015 4 [26] 551 Gabon -0.8683 12.6724 278 Anura Arthroleptidae Scotobleps gabonicus 2015 2 [26] 552 Gabon -0.8049 12.8124 255 Anura Arthroleptidae Scotobleps gabonicus 2015 2 [26] 553 Gabon -2.2377 13.5809 541 Anura Arthroleptidae Scotobleps gabonicus 2015 1 [26] 554 Gabon -0.226 12.298 213 Anura Bufonidae Sclerophrys camerunensis 2011 2 [23] 555 Gabon -0.039 12.291 254 Anura Bufonidae Sclerophrys camerunensis 2011 1 [23] this 556 Gabon 1.1544 12.6992 505 Anura Bufonidae Sclerophrys camerunensis 2013 1 study this 557 Gabon -1.8553 13.8542 407 Anura Bufonidae Sclerophrys gracilipes 2013 2 study 558 Gabon -0.6420 10.2174 110 Anura Bufonidae Sclerophrys gracilipes 2015 1 [26] 559 Gabon -0.6407 10.2187 40 Anura Bufonidae Sclerophrys regularis 2015 1 [26] 560 Gabon -0.1829 10.7768 54 Anura Bufonidae Sclerophrys regularis 2015 1 [26] 561 Gabon -0.8683 12.6724 278 Anura Bufonidae Sclerophrys regularis 2015 1 [26] 562 Gabon 0.5037 12.7941 532 Anura Conrauidae Conraua crassipes 2009 1 [2] 563 Gabon -0.6438 10.2207 110 Anura Conrauidae Conraua crassipes 2015 1 [26] 564 Gabon -0.6397 10.2132 70 Anura Conrauidae Conraua crassipes 2015 1 [26] 565 Gabon -0.0492 11.1643 174 Anura Conrauidae Conraua crassipes 2015 3 [26] 566 Gabon 0.5112 12.8028 506 Anura Hyperoliidae Afrixalus "quadrivittatus" 2009 1 [2] 567 Gabon -0.0595 11.1593 102 Anura Hyperoliidae Afrixalus dorsalis 2015 1 [26] 568 Gabon 0.5112 12.8028 506 Anura Hyperoliidae Afrixalus paradorsalis 2009 1 [2] 569 Gabon -2.1691 13.6395 479 Anura Hyperoliidae Cryptothylax greshoffii 2015 1 [26] 570 Gabon -2.3081 13.6099 531 Anura Hyperoliidae Hyperolius adspersus 2015 1 [26] this 571 Gabon 1.1544 12.6992 505 Anura Hyperoliidae Hyperolius cinnamomeoventris 2013 2 study 572 Gabon -0.1829 10.7768 54 Anura Hyperoliidae Hyperolius cinnamomeoventris 2015 2 [26] 573 Gabon -0.6420 10.2174 110 Anura Hyperoliidae Hyperolius cinnamomeoventris 2015 4 [26] 574 Gabon -0.1829 10.7768 54 Anura Hyperoliidae Hyperolius cinnamomeoventris 2015 1 [26] 575 Gabon -0.0595 11.1593 102 Anura Hyperoliidae Hyperolius cinnamomeoventris 2015 1 [26] 576 Gabon -2.1691 13.6395 479 Anura Hyperoliidae Hyperolius cinnamomeoventris 2015 1 [26] 577 Gabon -2.3081 13.5990 560 Anura Hyperoliidae Hyperolius concolor 2015 2 [26] 578 Gabon 0.4999 12.8018 493 Anura Hyperoliidae Hyperolius ocellatus 2009 1 [2] 579 Gabon 0.5037 12.7941 532 Anura Hyperoliidae Hyperolius ocellatus 2009 1 [2] 580 Gabon 0.5162 12.7946 532 Anura Hyperoliidae Hyperolius ocellatus 2009 1 [2] 581 Gabon 0.4536 10.2781 136 Anura Hyperoliidae Hyperolius ocellatus 2009 1 [2] 582 Gabon 0.294 12.566 499 Anura Hyperoliidae Hyperolius ocellatus 2010 2 [23] 583 Gabon -2.2381 13.5824 541 Anura Hyperoliidae Hyperolius ocellatus 2015 1 [26] 584 Gabon -0.6420 10.2174 110 Anura Hyperoliidae Hyperolius ocellatus 2015 2 [26] 585 Gabon -0.1829 10.7768 54 Anura Hyperoliidae Hyperolius ocellatus 2015 2 [26] 586 Gabon -0.8683 12.6724 278 Anura Hyperoliidae Hyperolius ocellatus 2015 1 [26] 587 Gabon -2.1577 13.6343 523 Anura Hyperoliidae Hyperolius ocellatus 2015 1 [26] 588 Gabon -2.1546 13.6414 523 Anura Hyperoliidae Hyperolius ocellatus 2015 2 [26] 589 Gabon -2.3197 13.6099 547 Anura Hyperoliidae Hyperolius pardalis 2015 1 [26] 590 Gabon -2.1245 13.6351 519 Anura Hyperoliidae Hyperolius pardalis 2015 1 [26] 591 Gabon -0.6420 10.2174 110 Anura Hyperoliidae Hyperolius phantasticus 2015 4 [26] 592 Gabon -0.6420 10.2174 110 Anura Hyperoliidae Hyperolius phantasticus 2015 1 [26] 593 Gabon -1.814 9.356 19 Anura Hyperoliidae Hyperolius sp. 2010 1 [23] 594 Gabon 0.6212 10.4076 496 Anura Hyperoliidae Hyperolius tuberculatus 2009 1 [2] this 595 Gabon 1.1544 12.6992 505 Anura Hyperoliidae Hyperolius tuberculatus 2013 2 study 596 Gabon -0.1848 10.7773 60 Anura Hyperoliidae Hyperolius tuberculatus 2015 1 [26] 597 Gabon -0.1829 10.7768 54 Anura Hyperoliidae Hyperolius tuberculatus 2015 3 [26] 598 Gabon -0.1848 10.7773 60 Anura Hyperoliidae Hyperolius tuberculatus 2015 1 [26] 599 Gabon -0.0521 11.1649 194 Anura Hyperoliidae Phlyctimantis leonardi 2015 1 [26] 600 Gabon 0.6256 10.3965 494 Anura Petropedetidae Petropedetes cf. vulpiae 2009 1 [2] 601 Gabon 0.4536 10.2781 136 Anura Petropedetidae Petropedetes palmipes 2009 2 [2] 602 Gabon 0.4455 10.2820 130 Anura Phrynobatrachidae Phrynobatrachus auritus 2009 1 [2] 603 Gabon 0.6256 10.3965 494 Anura Phrynobatrachidae Phrynobatrachus auritus 2009 2 [2] this 604 Gabon -0.3650 11.7906 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 605 Gabon -0.3649 11.7905 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 606 Gabon -0.3647 11.7906 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 607 Gabon -0.3642 11.7907 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 608 Gabon -0.3638 11.7907 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 609 Gabon -0.3638 11.7907 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 610 Gabon -0.3633 11.7903 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 2 study this 611 Gabon -0.3633 11.7902 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 612 Gabon -0.3626 11.7895 259 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 613 Gabon 1.1534 12.69895 505 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 614 Gabon 1.1536 12.699 505 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 615 Gabon 1.1538 12.6990 505 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 616 Gabon 1.1543 12.6990 505 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 617 Gabon 1.1550 12.6995 505 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 618 Gabon 1.1550 12.6995 505 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 619 Gabon 1.1555 12.6975 505 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 1 study this 620 Gabon -1.8553 13.8542 407 Anura Phrynobatrachidae Phrynobatrachus auritus 2013 6 study 621 Gabon -2.0250 13.9517 425 Anura Phrynobatrachidae Phrynobatrachus auritus 2018 1 [30] 622 Gabon -2.0183 13.9586 434 Anura Phrynobatrachidae Phrynobatrachus auritus 2018 1 [30] 623 Gabon -2.010 13.965 429 Anura Phrynobatrachidae Phrynobatrachus auritus 2018 1 [30] 624 Gabon -1.892 9.568 35 Anura Phrynobatrachidae Phrynobatrachus sp. 2010 2 [23] 625 Gabon -1.891 9.579 25 Anura Phrynobatrachidae Phrynobatrachus sp. 2010 1 [23] this 626 Gabon -1.6278 12.3631 752 Anura Phrynobatrachidae Phrynobatrachus sp. 2013 1 study 627 Gabon -0.1809 10.7783 54 Anura Pipidae Xenopus cf. parafraseri 2015 2 [26] 628 Gabon -0.6431 10.2182 110 Anura Pipidae Xenopus epitropicalis 2015 1 [26] this 629 Gabon -1.8553 13.8542 407 Anura Ptychadenidae Ptychadena "mascareniensis" D 2013 1 study this 630 Gabon 1.1544 12.6992 505 Anura Ptychadenidae Ptychadena aequiplicata 2013 1 study 631 Gabon -0.8683 12.6724 278 Anura Ptychadenidae Ptychadena perreti 2015 1 [26] 632 Gabon -1.814 9.356 19 Anura Ptychadenidae Ptychadena sp. 2010 1 [23] 633 Gabon 0.290 12.573 429 Anura Ptychadenidae Ptychadena sp. 2010 1 [23] 634 Gabon -0.039 12.291 254 Anura Ptychadenidae Ptychadena sp. 2011 2 [23] 635 Gabon 0.45358 10.27809 136 Anura Ptychadenidae Ptychadena sp. A 2009 1 [2] 636 Gabon 0.50367 12.79407 532 Anura Ptychadenidae Ptychadena sp. B 2009 1 [2] this 637 Gabon -1.8553 13.8542 407 Anura Ranidae Amnirana albolabris 2013 1 study this 638 Gabon -1.8553 13.8542 407 Anura Ranidae Amnirana albolabris 2013 1 study this 639 Gabon -0.3636 11.7872 259 Anura Ranidae Amnirana albolabris 2013 1 study this 640 Gabon -0.3636 11.7872 259 Anura Ranidae Amnirana albolabris 2013 1 study 641 Gabon -2.2381 13.5824 541 Anura Ranidae Amnirana albolabris 2015 1 [26] 642 Gabon -0.6420 10.2174 110 Anura Ranidae Amnirana albolabris 2015 1 [26] 643 Gabon -0.6407 10.2187 40 Anura Ranidae Amnirana albolabris 2015 1 [26] 644 Gabon -0.6420 10.2174 110 Anura Ranidae Amnirana albolabris 2015 5 [26] 645 Gabon -0.6407 10.2187 40 Anura Ranidae Amnirana albolabris 2015 1 [26] 646 Gabon -0.1772 10.7821 54 Anura Ranidae Amnirana albolabris 2015 1 [26] 647 Gabon -2.2397 13.5826 541 Anura Ranidae Amnirana albolabris 2015 1 [26] this 648 Gabon -1.6278 12.3631 752 Anura Ranidae Amnirana amnicola 2013 1 study this 649 Gabon 1.1544 12.6992 505 Anura Ranidae Amnirana amnicola 2013 1 study this 650 Gabon -1.8553 13.8542 407 Anura Ranidae Amnirana amnicola 2013 1 study this 651 Gabon 1.1544 12.6992 505 Anura Ranidae Amnirana lepus 2013 1 study 652 Gabon -2.2419 13.5875 553 Anura Ranidae Amnirana lepus 2015 1 [26] 653 Gabon 0.500 12.802 533 Anura Ranidae Amnirana sp. 2010 1 [23] 654 Gabon 0.506 12.796 532 Anura Ranidae Amnirana sp. 2010 1 [23] this 655 Gabon -1.8553 13.8542 407 Anura Ranidae Amnirana sp. 2013 1 study 656 Gabon 0.4999 12.8018 493 Anura Ranidae Amnirana sp. A 2009 1 [2] 657 Gabon 0.5112 12.8028 506 Anura Ranidae Amnirana sp. A 2009 1 [2] 658 Gabon 0.5162 12.7946 532 Anura Ranidae Amnirana sp. A 2009 1 [2] 659 Gabon 0.6256 10.3965 494 Anura Ranidae Amnirana sp. A 2009 1 [2] 660 Gabon 0.5112 12.8028 506 Anura Rhacophoridae Chiromantis rufescens 2009 1 [2] 661 Gabon 0.4536 10.2781 136 Anura Rhacophoridae Chiromantis rufescens 2009 1 [2] 662 Gabon -0.8683 12.6724 278 Anura Rhacophoridae Chiromantis rufescens 2015 1 [26] 663 Gabon -0.1848 10.7773 60 Anura Rhacophoridae Chiromantis rufescens 2015 1 [26] 664 Gabon -0.1750 10.7814 54 Anura Rhacophoridae Chiromantis rufescens 2015 1 [26] 665 Kenya -3.4055 38.3644 1401 Anura Bufonidae Sclerophrys gutturalis 2006 2 [27] 666 Kenya -1.3316 36.7990 1717 Anura Bufonidae Sclerophrys regularis 2006 1 [27] 667 Kenya -4.1805 39.4191 173 Anura Hyperoliidae Afrixalus fornasini 2006 1 [27] 668 Kenya -4.1805 39.4191 173 Anura Hyperoliidae Afrixalus sylvaticus 2006 1 [27] 669 Kenya 0.0431 36.3715 2323 Anura Hyperoliidae Hyperolius acuticeps 2006 10 [27] 670 Kenya -0.3684 36.8424 2197 Anura Hyperoliidae Hyperolius glandicolor 2006 21 [27] 671 Kenya -1.3316 36.7990 1717 Anura Hyperoliidae Hyperolius glandicolor 2006 17 [27] 672 Kenya -3.5050 38.3816 842 Anura Hyperoliidae Hyperolius glandicolor 2006 3 [27] 673 Kenya -3.4055 38.3644 1401 Anura Hyperoliidae Hyperolius glandicolor 2006 21 [27] 674 Kenya 0.0431 36.3715 2323 Anura Hyperoliidae Hyperolius glandicolor 2006 37 [27] 675 Kenya -1.1373 36.6789 2120 Anura Hyperoliidae Hyperolius glandicolor 2006 9 [27] 676‡ Kenya -3.4822 38.3406 1192 Anura Hyperoliidae Hyperolius glandicolor 2010 3 BdGPL [3] 677 Kenya 0.3528 34.8653 1605 Anura Hyperoliidae Hyperolius kivuensis 2006 11 [27] 678 Kenya 1.1063 35.1247 1869 Anura Hyperoliidae Hyperolius kivuensis 2006 9 [27] 679 Kenya 1.1063 35.1247 1869 Anura Hyperoliidae Hyperolius lateralis 2006 1 [27] 680‡ Kenya -4.2638 39.3975 294 Anura Hyperoliidae Hyperolius rubrovermiculatus 2010 1 BdGPL [3] 681 Kenya -4.1805 39.4191 173 Anura Hyperoliidae Hyperolius tuberilinguis 2006 3 [27] 682‡ Kenya -4.2638 39.3975 294 Anura Hyperoliidae Hyperolius tuberilinguis 2010 1 BdGPL [3] 683 Kenya 1.1063 35.1247 1869 Anura Hyperoliidae Hyperolius viridiflavus 2006 38 [27] 684 Kenya -1.3316 36.7990 1717 Anura Hyperoliidae Kassina senegalensis 2006 1 [27] 685 Kenya -3.5050 38.3816 842 Anura Phrynobatrachidae Phrynobatrachus acridoides 2006 1 [27] 686‡ Kenya -4.2638 39.3975 294 Anura Phrynobatrachidae Phrynobatrachus acridoides 2010 1 BdGPL [3] 687* Kenya 0.2944 35.3347 2212 Anura Pipidae Xenopus borealis 1934 1 [41] 688* Kenya 0.2944 35.3347 2212 Anura Pipidae Xenopus borealis 1934 1 [41] 689 Kenya -0.3684 36.8424 2197 Anura Pipidae Xenopus borealis 2006 3 [27] 690* Kenya -0.1000 35.1167 1240 Anura Pipidae Xenopus victorianus 1980 1 [41] 691 Kenya 0.3528 34.8653 1605 Anura Pipidae Xenopus victorianus 2006 1 [27] 692 Kenya -0.3684 36.8424 2197 Anura Ptychadenidae Ptychadena "mascareniensis" 2006 12 [27] 693 Kenya -1.3316 36.7990 1717 Anura Ptychadenidae Ptychadena "mascareniensis" 2006 6 [27] 694 Kenya -3.4055 38.3644 1401 Anura Ptychadenidae Ptychadena "mascareniensis" 2006 1 [27] 695 Kenya 1.0339 34.7695 2283 Anura Ptychadenidae Ptychadena anchietae 2006 5 [27] 696 Kenya -1.3316 36.7990 1717 Anura Ptychadenidae Ptychadena anchietae 2006 2 [27] 697 Kenya -4.1805 39.4191 173 Anura Ptychadenidae Ptychadena anchietae 2006 1 [27] 698 Kenya -3.4055 38.3644 1401 Anura Ptychadenidae Ptychadena anchietae 2006 4 [27] 699 Kenya -0.3684 36.8424 2197 Anura Pyxicephalidae Amietia angolensis 2006 7 [27] 700 Kenya 1.1063 35.1247 1869 Anura Pyxicephalidae Amietia angolensis 2006 14 [27] 701 Kenya -3.4055 38.3644 1401 Anura Pyxicephalidae Amietia angolensis 2006 6 [27] 702 Kenya -0.4114 36.7238 3118 Anura Pyxicephalidae Amietia wittei 2006 22 [27] 703† Lesotho -29.3440 28.5195 2032 Anura Pyxicephalidae Amietia delalandii 2004 5 [42] 704 Lesotho -29.5813 29.2887 2878 Anura Pyxicephalidae Amietia delalandii 2008-2012 [5] [38] 705† Lesotho -28.7433 28.7992 3034 Anura Pyxicephalidae Amietia hymenopus 2006 39 [36] 706† Lesotho -29.8604 27.6225 1514 Anura Pyxicephalidae Amietia vertebralis 2000 1 [42] 707† Lesotho -29.3021 28.5817 2076 Anura Pyxicephalidae Amietia vertebralis 2000 1 [42] 708† Lesotho -29.4398 29.3803 2795 Anura Pyxicephalidae Amietia vertebralis 2000 1 [42] 709† Lesotho -29.5815 29.2888 2878 Anura Pyxicephalidae Amietia vertebralis 2000 1 [42] 710† Lesotho -29.3440 28.5195 2032 Anura Pyxicephalidae Amietia vertebralis 2004 2 [42] 711 Lesotho -29.5813 29.2887 2878 Anura Pyxicephalidae Amietia vertebralis 2008-2012 - [38] 712 Malawi -15.8228 35.7198 1377 Anura Arthroleptidae Arthroleptis xenodactyloides 2009 2 [5] 713 Malawi -16.0240 35.5022 647 Anura Hyperoliidae Afrixalus crotalus 2009 1 [5] 714 Malawi -16.0178 35.5194 818 Anura Phrynobatrachidae Phrynobatrachus natalensis 2009 1 [5] 715 Malawi -15.8228 35.7198 1377 Anura Phrynobatrachidae Phrynobatrachus natalensis 2009 1 [5] 716* Malawi -13.9824 33.7829 1039 Anura Pipidae Xenopus laevis laevis 1969 1 [37] 717 Malawi -16.0240 35.5022 647 Anura Pipidae Xenopus muelleri 2009 1 [5] 718 Malawi -16.0178 35.5194 818 Anura Pyxicephalidae Amietia delalandii 2009 1 [5] 719 Malawi -15.9555 35.6880 1856 Anura Pyxicephalidae Amietia delalandii 2009 2 [5] 720 Malawi -15.9044 35.6744 2282 Anura Pyxicephalidae Amietia delalandii 2009 1 [5] 721 Malawi -15.9555 35.6880 1856 Anura Pyxicephalidae Amietia johnstoni 2009 2 [5] 722 Malawi -15.8925 35.6166 2047 Anura Pyxicephalidae Nothophryne broadleyi 2009 1 [5] 723 Malawi -15.9251 35.6775 2219 Anura Pyxicephalidae Strongylopus fuelleborni 2009 1 [5] 724 Morocco 35.5359 -5.3862 308 Anura Alytidae Discoglossus scovazzi 2006 1 [10] 725 Morocco 35.0439 -6.0462 43 Anura Alytidae Discoglossus scovazzi 2007 1 [10] 726 Morocco 35.0439 -6.0462 43 Anura Hylidae Hyla meridionalis 2007 1 [10] 727 Morocco 35.0381 -6.0292 43 Anura Pelobatidae Pelobates varaldii 2009 1 [10] 728 Mozambique -14.9524 38.2710 522 Anura Pipidae Xenopus muelleri 2013 1 [7] this 729 Namibia -24.2655 16.2395 1547 Anura Pyxicephalidae Amietia poyntoni 2007 5 study this 730 Namibia -24.2469 16.1036 1390 Anura Pyxicephalidae Amietia poyntoni 2007 7 study 731† Nigeria 6.3781 5.1725 52 Anura Arthroleptidae Arthroleptis sp. 1 2015 1 [4] Nigeria 6.3561 5.1980 41 Anura [4] 732 Nigeria 7.33 11.59 618 Anura Arthroleptidae Astylosternus sp. 2009 1 [33] 733 Nigeria 6.3 5.25 52 Anura Arthroleptidae Leptopelis hyloides 2007-2008 10 [25] 734 Nigeria 7.33 11.59 618 Anura Bufonidae Sclerophrys sp. 2009 1 [33] 735† Nigeria 6.3781 5.1725 52 Anura Dicroglossidae Hoplobatrachus occipitalis 2015 2 [4] Nigeria 6.3561 5.1980 41 Anura [4] 736 Nigeria 6.3 5.25 52 Anura Hyperoliidae Afrixalus dorsalis 2007-2008 74 [25] 737 Nigeria 6.3 5.25 52 Anura Hyperoliidae Afrixalus nigeriensis 2007-2008 35 [25] 738 Nigeria 6.3 5.25 52 Anura Hyperoliidae Afrixalus paradorsalis 2007-2008 4 [25] 739 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius concolor 2007-2008 2 [25] 740 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius fusciventris burtoni 2007-2008 4 [25] 741 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius picturatus 2007-2008 6 [25] 742 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius sp. 1 2007-2008 5 [25] 743 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius sp. 2 2007-2008 2 [25] 744 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius sp. 3 2007-2008 1 [25] 745 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius sp. 4 2007-2008 2 [25] 746 Nigeria 6.3 5.25 52 Anura Hyperoliidae Hyperolius sylvaticus 2007-2008 14 [25] 747 Nigeria 7.33 11.59 618 Anura Petropedetidae Petropedetes sp. 2009 1 [33] 748 Nigeria 6.3 5.25 52 Anura Phrynobatrachidae Phrynobatrachus calcaratus 2007-2008 12 [25] 749 Nigeria 6.3 5.25 52 Anura Phrynobatrachidae Phrynobatrachus liberiensis 2007-2008 12 [25] 750 Nigeria 6.3 5.25 52 Anura Phrynobatrachidae Phrynobatrachus plicatus 2007-2008 6 [25] 751 Nigeria 6.3 5.25 52 Anura Pipidae Xenopus tropicalis 2007-2008 88 [25] 752† Nigeria 6.3781 5.1725 52 Anura Ptychadenidae Ptychadena "mascareniensis" 2015 3 [4] Nigeria 6.3561 5.1980 41 Anura [4] 753† Nigeria 6.3781 5.1725 52 Anura Ptychadenidae Ptychadena bibroni 2015 1 [4] Nigeria 6.3561 5.1980 41 Anura [4] 754 Nigeria 6.3 5.25 52 Anura Ptychadenidae Ptychadena longirostris 2007-2008 1 [25] 755† Nigeria 6.3781 5.1725 52 Anura Ptychadenidae Ptychadena longirostris 2015 1 [4] Nigeria 6.3561 5.1980 41 Anura [4] 756 Nigeria 6.3 5.25 52 Anura Ptychadenidae Ptychadena pumilio 2007-2008 6 [25] 757† Nigeria 6.3781 5.1725 52 Anura Ptychadenidae Ptychadena pumilio 2015 1 [4] Nigeria 6.3561 5.1980 41 Anura [4] 758 Nigeria 6.3 5.25 52 Anura Ranidae Amnirana albolabris 2007-2008 3 [25] 759† Nigeria 6.3 5.25 52 Anura Rhacophoridae Chiromantis rufescens 2007-2008 1 [24] 760 Nigeria 6.3 5.25 52 Anura Rhacophoridae Chiromantis rufescens 2007-2008 40 [25] Rep. of the this 761 Congo -2.5422 13.5298 529 Anura Arthroleptidae Arthroleptis poecilonotus 2010 1 study Rep. of the this 762 Congo -2.6769 13.5872 637 Anura Arthroleptidae Arthroleptis poecilonotus 2010 1 study Rep. of the this 763 Congo -4.1875 14.9568 338 Anura Arthroleptidae Arthroleptis sylvaticus 2008 1 study Rep. of the this 764 Congo -2.5422 13.5298 529 Anura Arthroleptidae Arthroleptis sylvaticus 2010 1 study Rep. of the this 765 Congo -2.6769 13.5872 637 Anura Arthroleptidae Astylosternus batesi 2010 2 study Rep. of the this 766 Congo -2.3261 13.6621 512 Anura Arthroleptidae Leptopelis aubryioides 2010 1 study Rep. of the this 767 Congo 1.0840 17.3001 323 Anura Bufonidae Sclerophrys camerunensis 2008 5 study Rep. of the this 768 Congo 1.0840 17.3001 323 Anura Bufonidae Sclerophrys camerunensis 2008 1 study Rep. of the this 769 Congo 1.0840 17.3001 323 Anura Bufonidae Sclerophrys gracilipes 2008 1 study Rep. of the this 770 Congo 1.0840 17.3001 323 Anura Bufonidae Sclerophrys gracilipes 2008 1 study Rep. of the this 771 Congo 1.0840 17.3001 323 Anura Bufonidae Sclerophrys gracilipes 2008 1 study Rep. of the this 772 Congo -4.1875 14.9568 338 Anura Bufonidae Sclerophrys maculata 2008 1 study Rep. of the this 773 Congo -2.6769 13.5872 637 Anura Bufonidae Sclerophrys pusilla 2010 1 study Rep. of the this 774 Congo 1.0840 17.3001 323 Anura Bufonidae Sclerophrys regularis 2008 1 study Rep. of the this 775 Congo -4.1875 14.9568 338 Anura Bufonidae Sclerophrys regularis 2008 1 study Rep. of the this 776 Congo -4.5891 12.0557 130 Anura Bufonidae Sclerophrys regularis 2010 1 study Rep. of the this 777 Congo -4.1875 14.9568 323 Anura Bufonidae Sclerophrys sp. 2008 1 study Rep. of the this 778 Congo 1.0840 17.3001 338 Anura Bufonidae Sclerophrys sp. 2008 1 study Rep. of the this 779 Congo -2.5422 13.5298 529 Anura Bufonidae Sclerophrys tuberosus 2010 1 study Rep. of the this 780 Congo -3.9177 15.3952 712 Anura Hyperoliidae Afrixalus "quadrivittatus" 2008 2 study Rep. of the this 781 Congo -4.1875 14.9568 338 Anura Hyperoliidae Afrixalus osorioi 2008 1 study Rep. of the this 782 Congo -4.1875 14.9568 338 Anura Hyperoliidae Afrixalus sp. 2008 2 study Rep. of the this 783 Congo 1.0840 17.3001 323 Anura Hyperoliidae Cryptothylax greshoffii 2008 1 study Rep. of the this 784 Congo 1.0840 17.3001 323 Anura Hyperoliidae Hyperolius dartevellei 2008 1 study Rep. of the this 785 Congo 1.0840 17.3001 323 Anura Hyperoliidae Hyperolius dartevellei 2008 2 study Rep. of the this 786 Congo 1.0840 17.3001 323 Anura Hyperoliidae Hyperolius sp. 2008 1 study Rep. of the this 787 Congo -4.5891 12.0557 130 Anura Hyperoliidae Hyperolius sp. 2010 7 study Rep. of the this 788 Congo 1.0840 17.3001 323 Anura Phrynobatrachidae Phrynobatrachus cf. perpalmatus 2008 1 study Rep. of the this 789 Congo -4.1875 14.9568 338 Anura Phrynobatrachidae Phrynobatrachus cf. perpalmatus 2008 1 study Rep. of the this 790 Congo -4.1875 14.9568 338 Anura Phrynobatrachidae Phrynobatrachus sp. 2008 1 study Rep. of the this 791 Congo -4.1875 14.9568 338 Anura Phrynobatrachidae Phrynobatrachus sp. 2008 3 study Rep. of the this 792 Congo 1.0840 17.3001 323 Anura Pipidae Hymenochirus curtipes 2008 15 study Rep. of the this 793 Congo -2.3261 13.6621 512 Anura Pipidae Hymenochirus sp. 2010 1 study Rep. of the this 794 Congo 1.0840 17.3001 323 Anura Pipidae Xenopus cf. epitropicalis 2008 1 study Rep. of the this 795 Congo -4.1875 14.9568 338 Anura Pipidae Xenopus cf. epitropicalis 2008 1 study Rep. of the this 796 Congo -2.3261 13.6621 512 Anura Pipidae Xenopus pygmaeus 2010 4 study Rep. of the this 797 Congo -2.6769 13.5872 637 Anura Ranidae Amnirana albolabris 2010 1 study Rep. of the this 798 Congo -2.5422 13.5298 529 Anura Ranidae Amnirana amnicola 2010 1 study Rep. of the this 799 Congo -2.7929 13.5233 649 Anura Ranidae Amnirana amnicola 2010 1 study Rep. of the this 800 Congo -4.5891 12.0557 130 Anura Ranidae Amnirana sp. 2010 1 study 801 Rwanda -2.4772 29.1581 1949 Anura Arthroleptidae Arthroleptis sp. 2011 2 [35] 802 Rwanda -2.4768 29.1585 2051 Anura Hyperoliidae Afrixalus cf. laevis 2010 1 [35] 803 Rwanda -2.4772 29.1581 1949 Anura Hyperoliidae Afrixalus cf. laevis 2011 4 [35] 804 Rwanda -2.5455 28.9819 2040 Anura Hyperoliidae Hyperolius castaneus 2010 1 [35] 805 Rwanda -2.4810 29.1549 1949 Anura Hyperoliidae Hyperolius castaneus 2010 1 [35] 806 Rwanda -2.4846 29.1531 2055 Anura Hyperoliidae Hyperolius castaneus 2010 4 [35] 807 Rwanda -2.4810 29.1549 1949 Anura Hyperoliidae Hyperolius castaneus 2011 12 [35] 808 Rwanda -2.4556 29.2493 2629 Anura Hyperoliidae Hyperolius discodactylus 2010 4 [35] 809 Rwanda -2.5455 28.9819 2040 Anura Hyperoliidae Hyperolius kivuensis 2010 3 [35] 810 Rwanda -2.5453 28.9851 2029 Anura Hyperoliidae Hyperolius kivuensis 2010 1 [35] 811 Rwanda -2.4768 29.1585 2051 Anura Pyxicephalidae Amietia sp. 2010 1 [35] 812 Rwanda -2.4840 29.1531 2055 Anura Pyxicephalidae Amietia sp. 2011 2 [35] São Tomé 813* and Príncipe 1.589 7.380 619 Anura Arthroleptidae Leptopelis palmatus 2001 1 [23] São Tomé 814* and Príncipe 1.588 7.381 619 Anura Arthroleptidae Leptopelis palmatus 2006 3 [23] São Tomé 815* and Príncipe 1.622 7.385 246 Anura Arthroleptidae Leptopelis palmatus 2006 1 [23] São Tomé BdGPL 816‡ and Príncipe 0.288 6.603 1146 Anura Hyperoliidae Hyperolius molleri 2001 2 (1) [23], [3] São Tomé BdGPL 817‡ and Príncipe 0.298 6.730 70 Anura Hyperoliidae Hyperolius molleri 2001 2 (1) [23], [3] São Tomé 818* and Príncipe 0.300 6.732 68 Anura Hyperoliidae Hyperolius molleri 2001 4 [23] São Tomé 819* and Príncipe 0.319 6.738 8 Anura Hyperoliidae Hyperolius molleri 2001 1 [23] São Tomé 820* and Príncipe 1.644 7.398 169 Anura Hyperoliidae Hyperolius molleri 2001 2 [23] São Tomé 821* and Príncipe 1.652 7.416 180 Anura Hyperoliidae Hyperolius molleri 2001 1 [23] São Tomé 822 and Príncipe 0.306 6.619 879 Anura Hyperoliidae Hyperolius molleri 2012 3 [22] São Tomé 823 and Príncipe 0.308 6.617 879 Anura Hyperoliidae Hyperolius molleri 2012 2 [22] São Tomé 824 and Príncipe 0.288 6.613 1146 Anura Hyperoliidae Hyperolius molleri 2012 1 [22] São Tomé 825 and Príncipe 0.302 6.732 68 Anura Hyperoliidae Hyperolius molleri 2012 2 [22] São Tomé 826 and Príncipe 0.281 6.591 1398 Anura Hyperoliidae Hyperolius molleri 2012 2 [22] São Tomé 827 and Príncipe 0.296 6.638 760 Anura Hyperoliidae Hyperolius molleri 2012 1 [22] São Tomé 828 and Príncipe 0.288 6.624 1011 Anura Hyperoliidae Hyperolius molleri 2012 1 [22] São Tomé 829* and Príncipe 0.276 6.606 1238 Anura Hyperoliidae Hyperolius thomensis 2006 2 [23] São Tomé 830 and Príncipe 0.276 6.604 1240 Anura Hyperoliidae Hyperolius thomensis 2012 1 [22] São Tomé 831* and Príncipe 1.669 7.413 175 Anura Phrynobatrachidae Phrynobatrachus dispar 2001 1 [23] São Tomé 832* and Príncipe 1.639 7.396 177 Anura Phrynobatrachidae Phrynobatrachus dispar 2001 1 [23] São Tomé 833* and Príncipe 1.580 7.384 537 Anura Phrynobatrachidae Phrynobatrachus dispar 2006 1 [23] São Tomé 834* and Príncipe 1.588 7.381 619 Anura Phrynobatrachidae Phrynobatrachus dispar 2006 4 [23] São Tomé 835* and Príncipe 1.628 7.417 113 Anura Phrynobatrachidae Phrynobatrachus dispar 2006 1 [23] São Tomé 836* and Príncipe 0.333 6.729 19 Anura Phrynobatrachidae Phrynobatrachus leveleve 2001 1 [23] São Tomé 837* and Príncipe 0.302 6.552 467 Anura Phrynobatrachidae Phrynobatrachus leveleve 2001 1 [23] São Tomé 838* and Príncipe 0.254 6.645 399 Anura Phrynobatrachidae Phrynobatrachus leveleve 2006 5 [23] São Tomé 839* and Príncipe 0.316 6.549 539 Anura Phrynobatrachidae Phrynobatrachus leveleve 2011 1 [23] São Tomé 840* and Príncipe 0.333 6.729 19 Anura Ptychadenidae Ptychadena newtoni 2001 1 [23] São Tomé 841 and Príncipe 0.302 6.732 68 Anura Ptychadenidae Ptychadena newtoni 2012 1 [22] São Tomé BdGPL 842‡ and Príncipe 0.261 6.651 565 Gymnophiona Dermophiidae Schistometopum thomense 2001 11 (2) [23], [3] São Tomé 843* and Príncipe 0.275 6.653 510 Gymnophiona Dermophiidae Schistometopum thomense 2001 4 [23] São Tomé 844* and Príncipe 0.276 6.650 510 Gymnophiona Dermophiidae Schistometopum thomense 2006 1 [23] São Tomé 845* and Príncipe 0.247 6.466 11 Gymnophiona Dermophiidae Schistometopum thomense 2006 1 [23] São Tomé 846 and Príncipe 0.286 6.570 577 Gymnophiona Dermophiidae Schistometopum thomense 2012 1 [22] São Tomé 847* and Príncipe 0.254 6.630 399 Gymnophiona Dermophiidae Schistometopum thomense 2012 1 [23] 848 South Africa -29.43 29.91 1531 Anura Arthroleptidae Leptopelis xenodactylus 2008-2012 1 [38] 849 South Africa -29.0359 30.5619 1317 Anura Arthroleptidae Leptopelis xenodactylus 2008-2012 1 [38] 850 South Africa -29.0493 23.7686 989 Anura Bufonidae Sclerophrys poweri 2015-2016 1 [39] 851 South Africa -29.6831 17.8873 895 Anura Bufonidae Vandijkophrynus gariepensis 2015-2016 6 [39] 852† South Africa -29.6830 17.8874 895 Anura Bufonidae Vandijkophrynus robinsoni 2004 5 [42] 853 South Africa -29.6831 17.8873 895 Anura Bufonidae Vandijkophrynus robinsoni 2015-2016 2 [39] 854† South Africa -28.7133 28.9361 1511 Anura Heleophrynidae Hadromophryne natalensis 2006 5 [36] 855 South Africa -23.8165 30.0238 1181 Anura Heleophrynidae Hadromophryne natalensis 2008 12 [12] 856 South Africa -28.7491 28.8719 2865 Anura Heleophrynidae Hadromophryne natalensis 2008 3 [12] 857‡ South Africa -23.8165 30.0238 1181 Anura Heleophrynidae Hadromophryne natalensis 2008 1 BdCAPE [32] 858‡ South Africa Anura Heleophrynidae Hadromophryne natalensis 2009 1 BdCAPE [11] 859‡ South Africa -28.7199 28.9238 2273 Anura Heleophrynidae Hadromophryne natalensis 2010 1 BdCAPE [32] 860† South Africa -24.5902 30.8441 1177 Anura Heleophrynidae Hadromophryne natalensis 2013-2014 18 [9] 861† South Africa -24.5816 30.8712 1232 Anura Heleophrynidae Hadromophryne natalensis 2013-2014 3 [9] 862† South Africa -24.5522 30.8741 1816 Anura Heleophrynidae Hadromophryne natalensis 2013-2014 4 [9] 863† South Africa -24.5611 30.9114 1160 Anura Heleophrynidae Hadromophryne natalensis 2013-2014 1 [9] 864† South Africa -24.5831 30.8623 1405 Anura Heleophrynidae Hadromophryne natalensis 2013-2014 5 [9] 865† South Africa -24.5623 30.8713 1524 Anura Heleophrynidae Hadromophryne natalensis 2013-2014 1 [9] 866 South Africa -33.3251 25.3941 289 Anura Heleophrynidae Heleophryne hewitti 2008-2012 27 [38] 867 South Africa -33.9652 20.7044 383 Anura Heleophrynidae Heleophryne orientalis 2008-2012 2 [38] 868† South Africa -33.0892 19.0631 905 Anura Heleophrynidae Heleophryne purcelli 1998 3 [42] 869† South Africa -33.6823 22.2505 427 Anura Heleophrynidae Heleophryne regis 1981 1 [42] 870 South Africa -33.9821 18.4235 667 Anura Heleophrynidae Heleophryne rosei 2008-2012 17 [38] 871‡ South Africa -28.7198 28.9236 2273 Anura Heleophrynidae 2016 3 BdCAPE [32] 872 South Africa -29.56 30.25 1077 Anura Hyperoliidae Afrixalus spinifrons 2008-2012 [15] [38] 873 South Africa -29.99 30.9 27 Anura Hyperoliidae Afrixalus spinifrons 2008-2012 - [38] 874 South Africa -30.7919 30.7919 -9999 Anura Hyperoliidae Hyperolius pickersgilli 2008-2012 4 [38] 875 South Africa -28.7507 28.8682 3038 Anura Hyperoliidae Hyperolius pusillus 2008 3 [12] 876 South Africa -25.4270 30.9066 728 Anura Hyperoliidae Hyperolius viridiflavus 2008 2 [12] 877 South Africa -28.7507 28.8682 3038 Anura Hyperoliidae Hyperolius viridiflavus 2008 1 [12] 878† South Africa -33.6696 26.6461 1 Anura Hyperoliidae Kassina senegalensis 2004 1 [42] 879 South Africa -25.1071 31.0781 766 Anura Phrynobatrachidae Phrynobatrachus natalensis 2008 2 [12] 880 South Africa -25.4461 30.8848 686 Anura Phrynobatrachidae Phrynobatrachus natalensis 2008 1 [12] 881† South Africa -34.3206 18.4528 67 Anura Pipidae Xenopus gilli 1943 1 [42],[44] 882† South Africa -34.0706 18.5073 5 Anura Pipidae Xenopus gilli 1976 1 [42] 883* South Africa -34.3243 18.4501 67 Anura Pipidae Xenopus gilli 1982 2 [37] 884 South Africa -34.0706 18.5073 5 Anura Pipidae Xenopus laevis 1938 1 [42],[44] 885† South Africa -28.2861 29.1044 1640 Anura Pipidae Xenopus laevis 1972 1 [42] 886† South Africa -30.8405 28.5438 1531 Anura Pipidae Xenopus laevis 1972 1 [42] 887† South Africa -29.06 30.58 1065 Anura Pipidae Xenopus laevis 1973 1 [42] 888† South Africa -33.3460 20.0368 773 Anura Pipidae Xenopus laevis 1974 1 [42] 889† South Africa -28.5000 27.9168 1676 Anura Pipidae Xenopus laevis 1974 1 [42],[44] 890† South Africa -34.0357 18.5583 27 Anura Pipidae Xenopus laevis 1982 1 [42] 891† South Africa -27.5428 32.6752 -9999 Anura Pipidae Xenopus laevis 1985 1 [42] 892† South Africa -28.8067 26.1096 1269 Anura Pipidae Xenopus laevis 1987 1 [42] 893† South Africa -29.4127 25.0171 1171 Anura Pipidae Xenopus laevis 1991 1 [42] 894† South Africa -29.1554 26.5354 1337 Anura Pipidae Xenopus laevis 1991 1 [42] 895† South Africa -29.2089 30.9928 402 Anura Pipidae Xenopus laevis 1995 1 [42],[44] 896† South Africa -29.6830 17.8873 895 Anura Pipidae Xenopus laevis 1996 1 [42] 897† South Africa -29.3205 27.2815 1702 Anura Pipidae Xenopus laevis 1996 1 [42],[44] 898† South Africa -28.3482 24.8166 1174 Anura Pipidae Xenopus laevis 1998 1 [42],[44] 899‡ South Africa -34.5433 20.0441 70 Anura Pipidae Xenopus laevis 1999 [1] [29] 900‡ South Africa -34.1442 19.0242 310 Anura Pipidae Xenopus laevis 1999 [1] [29] 901‡ South Africa -33.4795 19.5916 523 Anura Pipidae Xenopus laevis 1999 [1] [29] 902‡ South Africa -34.0030 23.0391 243 Anura Pipidae Xenopus laevis 1999 [1] [29] 903† South Africa -33.8034 18.8669 169 Anura Pipidae Xenopus laevis 2001 17 [42] 904† South Africa -33.9266 18.8606 115 Anura Pipidae Xenopus laevis 2001 14 [42] 905† South Africa -34.0906 18.8414 25 Anura Pipidae Xenopus laevis 2002 17 [42] 906† South Africa -33.6336 19.0022 117 Anura Pipidae Xenopus laevis 2002 4 [42] 907† South Africa -34.2237 19.2072 86 Anura Pipidae Xenopus laevis 2002 14 [42] 908† South Africa -30.2042 17.9341 780 Anura Pipidae Xenopus laevis 2004 26 [42] 909‡ South Africa -29.9830 19.3743 956 Anura Pipidae Xenopus laevis 2005 1 BdGPL [34] 910 South Africa -25.6087 29.7121 1728 Anura Pipidae Xenopus laevis 2008 2 [12] 911 South Africa -27.9732 24.8282 1177 Anura Pipidae Xenopus laevis 2008 7 [12] 912 South Africa -27.9651 24.8290 1177 Anura Pipidae Xenopus laevis 2008 2 [12] 913 South Africa -27.7239 29.5239 1796 Anura Pipidae Xenopus laevis 2008 2 [12] 914‡ South Africa -22.9017 30.6961 492 Anura Pipidae Xenopus laevis 2008 1 BdGPL [32] 915 South Africa -30.2042 17.9336 780 Anura Pipidae Xenopus laevis 2008-2012 1 [38] 916 South Africa -30.2607 30.6109 439 Anura Ptychadenidae Ptychadena oxyrhynchus 2008 2 [12] 917‡ South Africa -27.6828 29.5786 1813 Anura Pyxicephalidae Amietia angolensis 2008 1 BdGPL [32] 918‡ South Africa -23.8500 30.0333 1308 Anura Pyxicephalidae Amietia angolensis 2008 1 BdGPL [32] 919‡ South Africa -29.0224 30.5811 1253 Anura Pyxicephalidae Amietia angolensis 2010 3 BdCAPE [32] 920† South Africa -24.8945 28.3320 1143 Anura Pyxicephalidae Amietia delalandii 2004 2 [42] 921† South Africa -28.6041 28.2245 1898 Anura Pyxicephalidae Amietia delalandii 2004 1 [42] 922† South Africa -26.9552 27.5923 1466 Anura Pyxicephalidae Amietia delalandii 2005-2006 1 [6] 923† South Africa -26.9281 27.1841 1317 Anura Pyxicephalidae Amietia delalandii 2005-2006 11 [6] 924† South Africa -26.8591 27.2884 1361 Anura Pyxicephalidae Amietia delalandii 2005-2006 6 [6] 925 South Africa -27.9732 24.8282 1177 Anura Pyxicephalidae Amietia delalandii 2008 2 [12] 926 South Africa -28.7632 28.9185 2768 Anura Pyxicephalidae Amietia delalandii 2008 1 [12] 927 South Africa -25.0139 30.8313 1342 Anura Pyxicephalidae Amietia delalandii 2008 3 [12] 928 South Africa -23.8165 30.0238 1181 Anura Pyxicephalidae Amietia delalandii 2008 12 [12] 929 South Africa -27.7239 29.5239 1796 Anura Pyxicephalidae Amietia delalandii 2008 10 [12] 930 South Africa -25.0856 30.7783 1005 Anura Pyxicephalidae Amietia delalandii 2008 1 [12] 931 South Africa -25.0894 30.7779 1141 Anura Pyxicephalidae Amietia delalandii 2008 2 [12] 932 South Africa -24.2800 28.5500 1600 Anura Pyxicephalidae Amietia delalandii 2008 3 [12] 933 South Africa -29.7732 30.8311 449 Anura Pyxicephalidae Amietia delalandii 2008-2012 1 [38] 934 South Africa -30.6864 26.7062 1306 Anura Pyxicephalidae Amietia delalandii 2015-2016 1 [39] 935 South Africa -29.0493 23.7686 989 Anura Pyxicephalidae Amietia delalandii 2015-2016 13 [39] 936 South Africa -28.8807 21.9856 855 Anura Pyxicephalidae Amietia delalandii 2015-2016 8 [39] 937 South Africa -28.4649 21.2434 798 Anura Pyxicephalidae Amietia delalandii 2015-2016 11 [39] 938† South Africa -32.5410 27.3663 932 Anura Pyxicephalidae Amietia fuscigula 2002 1 [28] 939† South Africa -32.5642 27.4299 827 Anura Pyxicephalidae Amietia fuscigula 2002 1 [42] 940† South Africa -32.3748 19.0621 594 Anura Pyxicephalidae Amietia fuscigula 2003 1 [21] 941† South Africa -29.6342 18.0069 1151 Anura Pyxicephalidae Amietia fuscigula 2003 10 [21] 942† South Africa -33.4198 22.2408 613 Anura Pyxicephalidae Amietia fuscigula 2003 2 [21] 943† South Africa -33.9986 19.0566 1075 Anura Pyxicephalidae Amietia fuscigula 2003 1 [21] 944† South Africa -33.0539 19.0796 695 Anura Pyxicephalidae Amietia fuscigula 2003 1 [21] 945† South Africa -32.3375 19.0247 350 Anura Pyxicephalidae Amietia fuscigula 2003 1 [21] 946† South Africa -29.6292 18.0342 1105 Anura Pyxicephalidae Amietia fuscigula 2003 8 [21] 947† South Africa -33.9333 18.8667 123 Anura Pyxicephalidae Amietia fuscigula 2003 1 [21] 948† South Africa -34.0099 20.4592 201 Anura Pyxicephalidae Amietia fuscigula 2003 3 [21] 949† South Africa -33.9483 18.4333 281 Anura Pyxicephalidae Amietia fuscigula 2003 3 [21] 950† South Africa -33.9479 18.4345 372 Anura Pyxicephalidae Amietia fuscigula 2003 3 [21] 951† South Africa -33.9652 20.7046 383 Anura Pyxicephalidae Amietia fuscigula 2003 1 [21] 952† South Africa -33.9531 25.5547 82 Anura Pyxicephalidae Amietia fuscigula 2004 1 [42] 953 South Africa -34.0909 18.4211 316 Anura Pyxicephalidae Amietia fuscigula 2008 3 [12] 954‡ South Africa -34.0933 18.4217 55 Anura Pyxicephalidae Amietia fuscigula 2008 2 BdGPL [32] 955 South Africa -32.5948 26.9468 1145 Anura Pyxicephalidae Amietia fuscigula 2008-2012 1 [38] 956‡ South Africa -34.0933 18.4217 55 Anura Pyxicephalidae Amietia fuscigula 2009 1 BdGPL [11] 957 South Africa -29.8001 24.4222 365 Anura Pyxicephalidae Amietia fuscigula 2015-2016 1 [39] 958 South Africa -28.7605 28.8981 2971 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 959 South Africa -28.7603 28.8973 2971 Anura Pyxicephalidae Amietia hymenopus 2015 3 [19] 960 South Africa -28.7556 28.8674 3038 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 961 South Africa -28.7526 28.8917 2979 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 962 South Africa -28.7525 28.8909 2993 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 963 South Africa -28.7504 28.8849 2993 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 964 South Africa -28.7502 28.8849 2993 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 965 South Africa -28.7501 28.8848 2993 Anura Pyxicephalidae Amietia hymenopus 2015 2 [19] 966 South Africa -28.7501 28.8846 2993 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 967 South Africa -28.7536 28.8884 2993 Anura Pyxicephalidae Amietia hymenopus 2015 1 [19] 968† South Africa -30.5520 17.9920 234 Anura Pyxicephalidae Amietia poyntoni 1996 5 [42] 969† South Africa -29.6967 18.0283 919 Anura Pyxicephalidae Amietia poyntoni 2004 6 [42] 970† South Africa -30.3148 18.0822 1401 Anura Pyxicephalidae Amietia poyntoni 2004 3 [42] 971† South Africa -30.2042 17.9341 780 Anura Pyxicephalidae Amietia poyntoni 2004 45 [42] 972† South Africa -29.6830 17.8874 895 Anura Pyxicephalidae Amietia poyntoni 2004 2 [42] 973 South Africa -25.6086 29.7121 1728 Anura Pyxicephalidae Amietia poyntoni 2008 1 [12] 974 South Africa -27.9732 24.8282 1177 Anura Pyxicephalidae Amietia poyntoni 2008 1 [12] 975 South Africa -27.7239 29.5239 1796 Anura Pyxicephalidae Amietia poyntoni 2008 2 [12] 976 South Africa -30.2042 17.9336 780 Anura Pyxicephalidae Amietia poyntoni 2008-2012 1 [38] 977 South Africa -28.7372 19.3045 1097 Anura Pyxicephalidae Amietia poyntoni 2015-2016 6 [39] 978 South Africa -28.9648 19.1463 342 Anura Pyxicephalidae Amietia poyntoni 2015-2016 15 [39] 979 South Africa -29.6831 17.8873 895 Anura Pyxicephalidae Amietia poyntoni 2015-2016 2 [39] 980 South Africa -25.0208 31.0117 587 Anura Pyxicephalidae Amietia sp. 2008 1 [12] 981† South Africa -28.7133 28.9361 1511 Anura Pyxicephalidae Amietia vertebralis 2006 [1] [36] 982 South Africa -28.7627 28.9179 2768 Anura Pyxicephalidae Amietia vertebralis 2008 12 [12] 983 South Africa -28.7419 28.9146 1909 Anura Pyxicephalidae Amietia vertebralis 2008 10 [12] 984 South Africa -28.7491 28.8719 2865 Anura Pyxicephalidae Amietia vertebralis 2008 3 [12] 985‡ South Africa -28.7627 28.9179 2994 Anura Pyxicephalidae Amietia vertebralis 2008 1 BdGPL [32] 986‡ South Africa -28.8446 29.0545 2072 Anura Pyxicephalidae Amietia vertebralis 2008 1 BdCAPE [32] 987 South Africa -33.7113 18.7231 80 Anura Pyxicephalidae Cacosternum aggestum 2008-2012 3 [38] 988† South Africa -33.6696 26.6461 1 Anura Pyxicephalidae Cacosternum boettgeri 2004 4 [42] 989 South Africa -27.7209 29.5216 1796 Anura Pyxicephalidae Cacosternum boettgeri 2008 1 [12] 990 South Africa -29.0359 30.5619 1317 Anura Pyxicephalidae Cacosternum striatum 2008-2012 2 [38] 991 South Africa -33.9964 18.4839 24 Anura Pyxicephalidae Microbatrachella capensis 2008-2012 5 [38] 992† South Africa -33.6696 26.6461 1 Anura Pyxicephalidae Strongylopus fasciatus 2004 3 [42] 993† South Africa -32.3375 19.0247 350 Anura Pyxicephalidae Strongylopus grayii 2003 1 [21] 994† South Africa -28.7133 28.9361 1511 Anura Pyxicephalidae Strongylopus grayii 2006 [1] [36] 995 South Africa -32.5948 26.9468 1145 Anura Pyxicephalidae Strongylopus grayii 2008-2012 2 [38] 996† South Africa -24.8945 28.3320 1143 Anura Pyxicephalidae Tomopterna cryptotis 2004 2 [42] 997 South Africa -29.0493 23.7686 989 Anura Pyxicephalidae Tomopterna cryptotis 2015-2016 1 [39] 998† South Africa -24.8945 28.3320 1143 Anura Pyxicephalidae Tomopterna natalensis 2004 1 [42] 999‡ South Africa -29.0224 30.5811 1253 Anura Pyxicephalidae 2010 1 BdCAPE [32] 1000‡ South Africa -30.5981 29.8945 839 Anura Pyxicephalidae 2015 1 BdGPL [32] 1001‡ South Africa -28.9136 19.0014 351 Anura Pyxicephalidae 2015 1 BdGPL [32] 1002‡ South Africa -28.6960 17.5974 208 Anura Pyxicephalidae 2015 1 BdGPL [32] 1003‡ South Africa -28.9136 19.0014 351 Anura Pyxicephalidae 2015 4 BdGPL [32] 1004‡ South Africa -28.7373 19.2992 416 Anura Pyxicephalidae 2015 2 BdGPL [32] 1005‡ South Africa -28.6960 17.5974 208 Anura Pyxicephalidae 2015 2 BdGPL [32] 1006‡ South Africa -32.5157 26.9346 1472 Anura Pyxicephalidae 2015 1 Hybrid [32] 1007‡ South Africa -30.7219 26.9065 1449 Anura Pyxicephalidae 2016 2 BdGPL [32] 1008‡ South Africa -28.7603 28.8964 2994 Anura Pyxicephalidae 2016 2 BdCAPE [32] 1009‡ South Africa -28.7506 28.8714 3124 Anura Pyxicephalidae 2016 2 BdCAPE [32] 1010‡ South Africa -28.7603 28.8964 2994 Anura Pyxicephalidae 2016 1 BdCAPE [32] 1011‡ South Africa -30.7219 26.9065 1449 Anura Pyxicephalidae 2016 1 Hybrid [32] 1012† Tanzania -8.5975 35.8381 1186 Anura Bufonidae Nectophrynoides asperginis 2003 [1] [43] 1013† Tanzania -8.515 35.865 1434 Anura Hyperoliidae Afrixalus sp. nov. 2005 5 [31] 1014† Tanzania -8.1703 36.0017 1569 Anura Hyperoliidae Afrixalus uluguruensis 2005 1 [31] 1015† Tanzania -7.8192 35.7789 1627 Anura Hyperoliidae Hyperolius acuticeps 2005 6 [31] 1016† Tanzania -8.5196 35.4611 1755 Anura Hyperoliidae Hyperolius pictus 2005 5 [31] 1017† Tanzania -8.5077 35.8608 1376 Anura Hyperoliidae Hyperolius pictus 2005 2 [31] 1018† Tanzania -8.5196 35.4611 1755 Anura Hyperoliidae Hyperolius pseudargus 2005 1 [31] 1019† Tanzania -8.1703 36.0017 1569 Anura Hyperoliidae Hyperolius puncticulatus 2005 2 [31] 1020† Tanzania -8.515 35.865 1434 Anura Hyperoliidae Hyperolius puncticulatus 2005 2 [31] 1021† Tanzania -7.7667 36.4250 1467 Anura Hyperoliidae Hyperolius puncticulatus 2005 2 [31] 1022† Tanzania -7.8250 36.3917 1710 Anura Hyperoliidae Hyperolius sp. 2005 1 [31] 1023† Tanzania -8.5196 35.4611 1755 Anura Hyperoliidae Hyperolius spinigularis 2005 1 [31] 1024† Tanzania -7.8192 35.7789 1627 Anura Hyperoliidae Kassina senegalensis 2005 4 [31] 1025† Tanzania -7.7686 36.9146 309 Anura Hyperoliidae Kassina senegalensis 2005 1 [31] 1026† Tanzania -8.4962 35.9424 293 Anura Microhylidae Phrynomantis bifasciatus 2005 2 [31] 1027† Tanzania -8.5864 35.8742 898 Anura Petropedetidae Arthroleptides yakusini 2003 [1] [43] 1028† Tanzania -8.5975 35.8381 1186 Anura Ptychadenidae Ptychadena aequiplicata 2003 [1] [43] 1029 Tanzania -3.1808 37.2511 1736 Anura Pyxicephalidae Amietia angolensis 2011 4 [45] 1030 Tanzania -3.0940 37.2657 2948 Anura Pyxicephalidae Amietia wittei 2011 3 [45] 1031 Tanzania -3.0967 37.2667 2601 Anura Pyxicephalidae Amietia wittei 2011 2 [45] 1032 Tanzania -3.2018 37.5175 2319 Anura Pyxicephalidae Amietia wittei 2011 1 [45] 1033 Tanzania -6.00 37.55 1237 Gymnophiona Herpelidae Boulengerula cf. uluguruensis 2008-2009 4 [15] 1034 Tanzania -5.49 37.49 1256 Gymnophiona Herpelidae Boulengerula cf. uluguruensis 2008-2009 1 [15] 1035 Tanzania -6.06 37.48 1596 Gymnophiona Scolecomorphidae Scolecomorphus cf. kirkii 2008-2009 1 [15] 1036 Tanzania -5.49 37.49 1256 Gymnophiona Scolecomorphidae Scolecomorphus cf. kirkii 2008-2009 1 [15] 1037 Uganda 1.724 31.528 1062 Anura Arthroleptidae Arthroleptis not reported 2013 1 [35] 1038 Uganda 0.5610 30.3578 1494 Anura Arthroleptidae Leptopelis christyi 2006 2 [13] 1039 Uganda 0.5610 30.3578 1494 Anura Arthroleptidae Leptopelis kivuensis 2006 9 [13] 1040 Uganda -0.9868 29.6351 1902 Anura Arthroleptidae Leptopelis kivuensis 2011 1 [35] 1041 Uganda -1.0195 29.7421 1992 Anura Arthroleptidae Leptopelis kivuensis 2011 2 [35] 1042 Uganda -1.0888 29.7542 2094 Anura Arthroleptidae Leptopelis kivuensis 2011 1 [35] 1043 Uganda -1.0863 29.7528 2094 Anura Arthroleptidae Leptopelis kivuensis 2011 1 [35] 1044 Uganda -1.0786 29.7461 2109 Anura Arthroleptidae Leptopelis kivuensis 2011 1 [35] 1045 Uganda -1.0504 29.7873 2237 Anura Arthroleptidae Leptopelis kivuensis 2011 3 [35] 1046 Uganda -1.0447 29.7782 2263 Anura Arthroleptidae Leptopelis kivuensis 2011 2 [35] 1047 Uganda -0.9913 29.6146 1535 Anura Arthroleptidae Leptopelis sp. 2011 1 [35] 1048 Uganda -0.9868 29.6351 1902 Anura Arthroleptidae Leptopelis sp. 2011 1 [35] 1049 Uganda 0.5610 30.3578 1494 Anura Bufonidae Sclerophrys funerea 2006 3 [13] 1050‡ Uganda 1.7271 31.5469 1077 Anura Bufonidae Sclerophrys sp. 2012 1 BdGPL [32] 1051 Uganda -1.0888 29.7542 2094 Anura Hyperoliidae Hyperolius castaneus 2011 1 [35] 1052 Uganda -1.1193 29.7082 1846 Anura Hyperoliidae Hyperolius castaneus 2011 1 [35] 1053 Uganda -1.0976 29.6997 2259 Anura Hyperoliidae Hyperolius castaneus 2011 1 [35] 1054 Uganda -0.9913 29.6146 1535 Anura Hyperoliidae Hyperolius cinnamomeoventris 2011 1 [35] 1055 Uganda -0.9760 29.6869 1588 Anura Hyperoliidae Hyperolius cinnamomeoventris 2011 1 [35] 1056 Uganda 0.5610 30.3578 1494 Anura Hyperoliidae Hyperolius kivuensis 2006 3 [13] 1057* Uganda 1.725 31.547 1102 Anura Hyperoliidae Hyperolius not reported 2013 1 [35] 1058 Uganda -0.9868 29.6351 1902 Anura Hyperoliidae Hyperolius sp. 6 2011 1 [35] 1059 Uganda -0.9913 29.6146 1535 Anura Hyperoliidae Hyperolius sp. X 2011 2 [35] 1060 Uganda -1.0786 29.7461 2109 Anura Hyperoliidae Hyperolius sp. X 2011 1 [35] 1061 Uganda -0.9868 29.6351 1902 Anura Hyperoliidae Hyperolius viridiflavus pitmani 2011 2 [35] 1062* Uganda 1.724 31.528 1062 Anura Phrynobatrachidae Phrynobatrachus not reported 2013 1 [35] 1063 Uganda -0.9868 29.6351 1902 Anura Pipidae Xenopus cf. laevis 2011 1 [35] 1064 Uganda -1.1197 29.7060 1846 Anura Pipidae Xenopus cf. wittei 2011 1 [35] 1065 Uganda -1.0976 29.6997 2259 Anura Pipidae Xenopus cf. wittei 2011 1 [35] 1066* Uganda -1.2303 29.8210 2131 Anura Pipidae Xenopus victorianus 1934 2 [37] 1067* Uganda 0.567 30.350 1483 Anura Pipidae Xenopus victorianus 1994 1 [41] 1068* Uganda 0.567 30.350 1483 Anura Pipidae Xenopus victorianus 1994 1 [41] 1069 Uganda 0.5610 30.3578 1494 Anura Pipidae Xenopus wittei 2006 5 [13] 1070 Uganda 0.5610 30.3578 1494 Anura Ptychadenidae Ptychadena "mascareniensis" 2006 2 [13] 1071 Uganda -0.8956 29.7299 1438 Anura Ptychadenidae Ptychadena achietae 2011 1 [35] 1072 Uganda -1.0997 29.7924 2084 Anura Ptychadenidae Ptychadena chrysogaster 2011 1 [35] 1073 Uganda -0.9771 29.6881 1588 Anura Ptychadenidae Ptychadena chrysogaster 2011 1 [35] 1074 Uganda -0.9868 29.6351 1902 Anura Pyxicephalidae Amietia angolensis 2011 1 [35] 1075 Uganda -1.0776 29.6445 1876 Anura Pyxicephalidae Amietia angolensis 2011 1 [35] 1076 Uganda -1.0343 29.7709 2163 Anura Pyxicephalidae Amietia angolensis 2011 1 [35] 1077† Uganda 0.3599 30.0229 1834 Anura Pyxicephalidae Amietia ruwenzorica 2005 6 [40] 1078 Uganda -0.9913 29.6146 1535 Anura Pyxicephalidae Amietia sp. 1 2011 1 [35] 1079 Zimbabwe -21.9816 29.9360 507 Anura Hyperoliidae Hyperolius sp. 2008 1 [12] 1080 Zimbabwe -21.9816 29.9360 507 Anura Hyperoliidae Hyperolius viridiflavus 2008 3 [12] 1081 Zimbabwe -21.9816 29.9360 507 Anura Hyperoliidae Kassina senegalensis 2008 4 [12] 1082 Zimbabwe -21.9816 29.9360 507 Anura Pipidae Xenopus laevis 2008 7 [12] 1083 Zimbabwe -21.9816 29.9360 507 Anura Pyxicephalidae Amietia delalandii 2008 5 [12] 1084 Zimbabwe -21.9816 29.9360 507 Anura Pyxicephalidae Cacosternum boettgeri 2008 31 [12]

References 1. Baláž, V., O. Kopecký, and V. Gvoždík V. 2012. Presence of the amphibian chytrid pathogen confirmed in Cameroon. Herpetological Journal 22:191–194. 2. Bell, R.C., A.V. Gata Garcia, B.L. Stuart, and K.R. Zamudio. 2011. High prevalence of the amphibian chytrid pathogen in Gabon. EcoHealth 8:116–120. 3. Byrne, A.Q., V.T. Vredenburg, A. Martel, . . . and E.B. Rosenblum. 2019. Cryptic diversity of a widespread global pathogen reveals new threats for amphibian conservation. Proceedings of the National Academy of Sciences of the United States of America 116:20382–20387. 4. Chinemerem, I.G. 2017. A survey of chytridiomycosis among amphibians (anurans) in Okomu Rubber Plantation, Edo State Nigera. Department of Animal and Environmental Biology, Faculty of Life Sciences, University of Benin, Benin City, Nigeria. 5. Conradie W., J. Harvey J, A. Kotzé, D.L. Dalton, and M.J. Cunningham. 2011a. Confirmed amphibian chytrid in mount Mulanje Area, Malawi. Herpetological Review 42:369–371. 6. Conradie, W., C. Weldon, K.G. Smith, L.H. du Preez. 2011b. Seasonal pattern of chytridiomycosis in common rover frog (Amietia angolensis) tadpoles in the South African Grassland Biome. African Zoology 46:95–102. 7. Conradie, W., G.B. Bittencourt-Silva, S.P. Loader, D.L. Dalton, and K.A. Tolley. 2016. Batrachochytrium dendrobatidis Survey of Amphibians in the Northern Mozambique “Sky Islands” and Low-lying Areas. Herpetological Review 47:42–46. 8. Doherty-Bone, T.M., N.L. Gonwouo, M. Hirschfeld, . . . and A.A. Cunningham. 2013. Batrachochytrium dendrobatidis in amphibians of Cameroon, including first records of infected caecilian hosts. Diseases of Aquatic Organisms 102:187–194. 9. DuToit, K.H. 2016. Factors affecting the occurrence and abundance of the Natal Cascade Frog, Hadromophryne natalensis, at Mariepskop, South Africa. M.Sc dissertation, University of Pretoria, South Africa. 10. El Mouden, E.H., T. Slimani, D. Donaire, S. Fernández-Beaskoetxea, M.C. Fisher, and J. Bosch. 2011. First Record of the chytrid fungus Batrachochytrium dendrobatidis in North Africa. Herpetological Review 42:71–75. 11. Farrer, R.A., L.A. Weinert, J. Bielby, . . . and M.C. Fisher. 2011. Multiple emergences of genetically diverse amphibian-infecting chytrids include a globalized hypervirulent recombinant lineage. Proceedings of the National Academy of Sciences of the United States of America 108:18732–18736. 12. Gericke, M.C. 2008. Aspects of Amphibian Chytrid Infections in South Africa. M.Sc dissertation, North-West University. 13. Goldberg, T.L., A.M. Readel, and M.H. Lee. 2007. Chytrid fungus in frogs from an equatorial African montane forest in western Uganda. Journal of Wildlife Diseases 43:521–524. 14. Gower, D.J., T.M. Doherty-Bone, R.K. Aberra, A. Mengistu, S. Schwaller, M. Menegon, R.O. de Sá, S.A. Saber, A.A. Cunningham, and S.P. Loader. 2012. High prevalence of the amphibian chytrid fungus (Batrachochytrium dendrobatidis) across multiple taxa and localities in the highlands of Ethiopia. Herpetological Journal 22:225–233. 15. Gower, D.J., T. Doherty-Bone, S.P. Loader, M. Wilkinson, M.T. Kouete, B. Tapley B, et al. 2013. Batrachochytrium dendrobatidis Infection and Lethal Chytridiomycosis in Caecilian Amphibians (Gymnophiona). Ecohealth 10:173–183. 16. Greenbaum, E., C. Kusamba, M.M. Aristote, and K. Reed. 2008. Amphibian chytrid fungus infections in Hyperolius (Anura: Hyperoliidae) from eastern Democratic Republic of Congo. Herpetological Review 39:70–73. 17. Greenbaum, E., J. Meece, K. Reed, and C. Kusamba. 2014. Amphibian chytrid infections in non-forested habitats of Katanga, Democratic Republic of the Congo. Herpetological Review 45:610–614. 18. Greenbaum, E., J. Meece, K.D. Reed, and C. Kusamba. 2015. Extensive occurrence of the amphibian chytrid fungus in the Albertine Rift, a Central African amphibian hotspot. Herpetological Journal 25:91–100. 19. Griffiths, S.M., X.A. Harrison, C. Weldon. M.D. Wood, A. Pretorius, K. Hopkins, G. Fox, R.F. Preziosi, and R.E. Antwis. 2018. Genetic variability and ontogeny predict microbiome structure in a disease-challenged montane amphibian. The ISME Journal 12:2506–2517. 20. Hirschfeld, M., D.C. Blackburn, T.M. Doherty-Bone, L.N. Gonwouo, S. Ghose, and M-O. Rödel. 2016. Dramatic declines of montane frogs in a Central African biodiversity hotspot. PLoS ONE 11:e0155129. 21. Hopkins, S., and A. Channing A. 2003. Chytrid fungus in northern and western Cape frog populations, South Africa. Herpetological Review 34:334–336. 22. Hydeman, M.E., R.C. Bell, R.C. Drewes, and K.R. Zamudio. 2013. Amphibian chytrid fungus confirmed in endemic frogs and caecilians on the Island of São Tomé, Africa. Herpetological Review 44(2):254–7. 23. Hydeman, M.E., A.V. Longo, G. Velo-Antón, D. Rodriguez, K.R. Zamudio, and R.C. Bell. 2017. Prevalence and genetic diversity of Batrachochytrium dendrobatidis in Central African island and continental amphibian communities. Ecology and Evolution 7:7729–38. 24. Imasuen, A.A., M.S.O. Aisen, C. Weldon, D.L. Dalton DL, A. Kotze A, and LH. du Preez. 2011. Occurrence of Batrachochytrium dendrobatidis in Amphibian Populations of Okomu National Park, Nigeria. Herpetological Review 42:379–382. 25. Imasuen, A.A., C. Weldon, M.S.O. Aisien, and L. H. Du Preez. 2009. Amphibian chytridiomycosis: First report in Nigeria from the skin of Chiromantis rufescens. Froglog 90:6–8. 26. Jongsma, G.F.M., A.B. Kaya, J.-A. Yoga, . . . D.C. Blackburn. 2016. Widespread presence and high prevalence of Batrachochytrium dendrobatidis in Gabon. Herpetological Review 47:227–230. 27. Kielgast, J., D. Rödder, M. Veith, and S. Lötters. 2010. Widespread occurrence of the amphibian chytrid fungus in Kenya. Animal Conservation 13:1–8. 28. Lane, E., C. Weldon, and J. Bingham. 2003. Histological evidence of chytridiomycosis in a free-ranging amphibian (Afrana fuscigula (Anura: Ranidae)) in South Africa. Journal of the South African Veterinary Association 74:20–21. 29. Mendez, D., R. Speare, A.A. Cunningham. 1999. In: Speare R, Berger L. (2000) Global distribution of chytridiomycosis in amphibians. Available: http://www.jcu.edu.au/school/phtm/PHTM/frogs/chyglob.htm. Accessed on 26 April 2011. 30. Miller, C.A., G.C. Tasse Taboue, M.M.P. Ekane, M. Robak, P.R. Sesink Clee, C. Richards-Zawacki, E.B. Fokam, N.A. Fuashi, and N.M. Anthony. 2018. Distribution modeling and lineage diversity of the chytrid fungus Batrachochytrium dendrobatidis (Bd) in a central African amphibian hotspot. PLoS ONE 13:e0199288. 31. Moyer, D., and C. Weldon. 2006. Chytrid distribution and pathogenicity among frogs of the Udzungwa Mountains Tanzania. Final report. CEPF Small Grant Final Project Completion. March 2006. Available at: https://www.cepf.net/sites/default/files/chytrid-distribution- udzungwa-mountains-report.pdf. 32. O’Hanlon, S.J., A. Rieux, R.A., Farrer, . . . M.C. Fisher. 2018. Recent Asian origin of chytrid fungi causing global amphibian declines. Science 627:621–627. 33. Reeder, N.M.M., T.L. Cheng, V.T. Vredenburg, D.C. Blackburn. 2011. Survey of the chytrid fungus Batrachochytrium dendrobatidis from montane and lowland frogs in eastern Nigeria. Herpetology Notes 4:83–86. 34. Rosenblum, E.B., T.Y. James, K.R. Zamudio, T.J. Poorten. D. Ilut, D. Rodriguez, J.M. Eastman, K. Ricjards-Hrdlicka, S. Joneson, T.S. Jenkinson, J.E. Longcore, G.P. Olea, L.F. Toledo, M.L. Arellano. E.M. Medina, S. Restrepo, S.V. Flechas, L. Berger, C.J. Briggs and J.E. Stajich. 2013. Complex history of the amphibian-killing chytrid fungus revealed with genome resequencing data. Proceedings of the National Academy of Sciences of the United States of America 110:9385–9390. 35. Seimon, T.A., S. Ayebare, R. Sekisambu, . . . and A.J. Plumptre. 2015. Assessing the threat of amphibian chytrid fungus in the Albertine Rift: Past, present and future. PLoS ONE 10: e0145841. 36. Smith, K.G., C. Weldon, W. Conradie and L.H. du Preez. 2007. Relationships among size, development, and Batrachochytrium dendrobatidis infection in African tadpoles. Diseases of Aquatic Organisms 74:159–164. 37. Soto-Azat, C., B.T. Clarke, J.C. Poynton and A.A. Cunningham. 2010. Widespread historical presence of Batrachochytridium dendrobatidis in African pipid frogs. Diversity and Distributions 16:126–131. 38. Tarrant, J. 2012. Conservation Assessment of Threatened Amphibians in KwaZulu-Natal, and a National Assessment of Chytrid Infection in South African Threatened Species. Ph.D. Dissertation, North-West University. 39. Verster, J.R. 2018. Spatial Epidemiology of Amphibian Chytridiomycosis in the Orange River system of South Africa. M.Sc dissertation, North-West University, 79 pp. 40. Viertel, B., M. Veith, S. Schick, A. Channing, S. Kigoolo, O. Baeza-Urrea, U. Sinsch, and S. Lötters. 2012. The stream-dwelling larva of the Ruwenzori river frog, Amietia ruwenzorica, its buccal cavity and pathology of chytridiomycosis. Zootaxa 3400:43–57. 41. Vredenburg, V.T., S.A. Felt, E.C. Morgan, S.V.G. McNally, S. Wilson & S.L. Green. 2013. Prevalence of Batrachochytrium dendrobatidis in Xenopus collected in Africa (1871–2000) and in California (2001–2010). PLoS ONE 8:e63791. 42. Weldon, C. 2005. Chytridiomycosis, an Emerging Infectious Disease of Amphibians in South Africa. Ph.D. dissertation, North-West University, South Africa. 43. Weldon, C. and L. Du Preez. 2004. Decline of the Kihansi spray toad, Nectophrynoides asperginis, from the Udzungwa mountains, Tanzania. Froglog 62:2–3. 44. Weldon, C., du Preez L.H., Hyatt A.D., Muller R., and Speare R. (2004) Origin of the amphibian chytrid fungus. Emerg Infect Diseases 10:2100–2105. 45. Zancolli, G., A. Storfer, and M.-O. Rödel. 2013. Detection of Batrachochytrium dendrobatidis in river frogs (genus Amietia) on Mount Kilimanjaro, Tanzania. Herpetological Review 44:611–614. 46. Frost, D.R. 2020. Amphibian Species of the World: an Online Reference. Version 6.0. Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/index.html. American Museum of Natural History, New York, USA. Accessed on 7 January 2020. SUPPLEMENTAL MATERIAL S2.—Environmental parameters used in environmental niche modelling (ENM) with a short description of the parameter and the source of the original data. ENM Variable Explanation Data source

Rain high Highest precipitation value; derived from the averages per month from 1950-2000 [1]

Rain low Lowest precipitation value; derived from the averages per month from 1950-2000 [1]

Rain SD Standard deviation of precipitation; derived from the averages per month from 1950-2000 [1]

Rain total Total annual precipitation; derived from the averages per month from 1950-2000 [1]

Temp. min low Lowest value of the minimum temperatures; derived from the averages per month from [1] 1950-2000

Temp. min high Highest value of the minimum temperatures; derived from the averages per month from [1] 1950-2000

Temp. min SD Standard deviation of the minimum temperatures; derived from the averages per month [1] from 1950-2000

Temp. max low Lowest value of the maximum temperatures; derived from the averages per month from [1] 1950-2000

Temp. max high Highest value of the maximum temperatures; derived from the averages per month from [1] 1950-2000

Temp. max SD Standard deviation of the maximum temperatures; derived from the averages per month [1] from 1950-2000

SPOT4 2 Annual average of vegetation derived from the wavelength 0.78–0.89µm of the SPOT4 [2] satellite for year 2000

SPOT4 3 Annual average vegetation derived from the wavelength 0.62–0.68µm of the SPOT4 [2] satellite for year 2000

MODIS bare Average percentage of bare ground from monthly composites from all 7 band lengths [3] derived from the MODIS satellite for year 2001

MODIS herb Average percentage of herbaceous cover from monthly composites from all 7 band lengths [4] derived from the MODIS satellite for year 2001

MODIS tree Average percentage of tree cover from monthly composites from all 7 band lengths derived [5] from the MODIS satellite for year 2001

Elev. var. ln-transformed and multiplied by 10 variation in elevation via 9x9 moving window derived [6] from SRTM30 data

Elev. contr. ln-transformed and multiplied by 10 contrast in elevation via 3x3 moving window derived [6] from SRTM30 data

References 1. Hijmans, R.J., S.E. Cameron, J.L. Parra, P.G. Jones, and A. Jarvis. 2005. Very high resolution interpolated climate surfaces for global land areas. Internationat Journal of Climatology 25:1965– 1978. 2. Arnaud, M., and M. Leroy M. 1991. SPOT 4: a new generation of SPOT satellites. ISPRS J Photogramm 46:205–215. 3. Hansen, M., R. DeFries, J.R. Townshend, M. Carroll, C. Dimiceli, et al. 2003a. Vegetation Continuous Fields MOD44B, 2001 Percent bare ground cover, Collection 3, Maryland, University of Maryland, College Park. 4. Hansen, M., R. DeFries, J.R. Townshend, M. Carroll, C. Dimiceli, et al. 2003b. Vegetation Continuous Fields MOD44B, 2001 Percent herbaceous ground cover, Collection 3, Maryland, University of Maryland, College Park. 5. Hansen, M., R. DeFries, J.R. Townshend, M. Carroll, C. Dimiceli, et al. 2003c. Vegetation Continuous Fields MOD44B, 2001 Percent tree cover, Collection Maryland, University of Maryland, College Park. 6. Farr, T.G., P.A. Rosen, E. Caro, . . . D. Alsdorf. 2007. The shuttle radar topography mission. Reviews of Geophysics 45:1–33. SUPPLEMENTAL MATERIAL S3.—Positive Bd results for newly sampled sites in Cameroon (including Cameroon-Nigeria border), Democratic Republic of Congo, Gabon, Namibia, and Republic of Congo.

Bd Intensity Country Site Lat Long Family Genus Species Year Specimen Voucher (GE) Adamaoua: Tchabal Mbabo, 1 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_110 0.698 Adamaoua: Tchabal Mbabo, 2 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_371 19.500 Adamaoua: Tchabal Mbabo, 3 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_372 20.450 Adamaoua: Tchabal Mbabo, 4 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_373 374.500 Adamaoua: Tchabal Mbabo, 5 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_374 468.000 Adamaoua: Tchabal Mbabo, 6 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_375 569.500 Adamaoua: Tchabal Mbabo, 7 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_376 78.300 Adamaoua: Tchabal Mbabo, 8 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_109 8.075 Adamaoua: Tchabal Mbabo, 9 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_111 7.155 Adamaoua: Tchabal Mbabo, 10 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_128 2.410 Adamaoua: Tchabal Mbabo, 11 Cameroon Foungoy 7.24570 12.07280 Pipidae Xenopus sp. nov. 2016 vgCM16_129 5.185 Adamaoua: Tchabal Mbabo, 12 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_084 ######## Adamaoua: Tchabal Mbabo, 13 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_085 3545.000 Adamaoua: Tchabal Mbabo, 14 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_086 3295.000 Adamaoua: Tchabal Mbabo, 15 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_087 224.500 Adamaoua: Tchabal Mbabo, 16 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_088 12.050 Adamaoua: Tchabal Mbabo, 17 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_089 330.000 Adamaoua: Tchabal Mbabo, 18 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_090 305.000 Adamaoua: Tchabal Mbabo, 19 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_091 52.900 Adamaoua: Tchabal Mbabo, 20 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_092 4565.000 Adamaoua: Tchabal Mbabo, 21 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_093 28.100 Adamaoua: Tchabal Mbabo, 22 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_094 43.500 Adamaoua: Tchabal Mbabo, 23 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_105 164.000 Adamaoua: Tchabal Mbabo, 24 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_107 3.015 Adamaoua: Tchabal Mbabo, 25 Cameroon Foungoy 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_108 96.550

Adamaoua: Tchabal Mbabo, 26 Cameroon Foungoy, camp 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_117 43.900 Adamaoua: Tchabal Mbabo, 27 Cameroon Foungoy, camp 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_118 1.171

Adamaoua: Tchabal Mbabo, 28 Cameroon Foungoy, camp 7.25220 12.05920 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_126 33.800 Adamaoua: Tchabal Mbabo, Foungoy, 29 Cameroon stream 7.25750 12.05250 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_063 5460.000 Adamaoua: Tchabal Mbabo, Foungoy, stream in 30 Cameroon pasture 7.24700 12.07200 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_101 56.650 Adamaoua: Tchabal Mbabo, Foungoy, 31 Cameroon watering place 7.26350 12.05160 Arthroleptidae Leptopelis nordequatorialis 2016 vgCM16_120 3.135 Babungo, Bamenda 32 Cameroon Highlands 6.04880 10.42585 Hyperoliidae Afrixalus quadrivittatus 2009 NMP:P6V 74545 3750.000 Babungo, Bamenda 33 Cameroon Highlands 6.04880 10.42585 Phrynobatrachidae Phrynobatrachus jimzimkusi 2009 NMP:P6V 74525/1 2.065 Babungo, Bamenda 34 Cameroon Highlands 6.04880 10.42585 Phrynobatrachidae Phrynobatrachus jimzimkusi 2009 NMP:P6V 74525/3 5.925 Babungo, Bamenda 35 Cameroon Highlands 6.04880 10.42585 Phrynobatrachidae Phrynobatrachus jimzimkusi 2009 NMP:P6V 74525/4 622.000 Babungo, Bamenda 36 Cameroon Highlands 6.04880 10.42585 Phrynobatrachidae Phrynobatrachus jimzimkusi 2009 NMP:P6V 74525/5 1.845 Babungo, Bamenda 37 Cameroon Highlands 6.04880 10.42585 Phrynobatrachidae Phrynobatrachus jimzimkusi 2009 NMP:P6V 74525/6 239.500 Babungo, Bamenda 38 Cameroon Highlands 6.04880 10.42585 Phrynobatrachidae Phrynobatrachus jimzimkusi 2009 NMP:P6V 74525/7 2345.000 Babungo, Bamenda 39 Cameroon Highlands 6.04880 10.42585 Pipidae Xenopus eysoole 2009 NMP:P6V 74555 4.745 40 Cameroon Banyo 6.73184 11.82868 Arthroleptidae Astylosternus diadematus 2009 NMP:P6V 74587/2 0.652 41 Cameroon Banyo 6.73184 11.82868 Arthroleptidae Leptopelis notatus 2009 NMP:P6V 74600/2 0.190 42 Cameroon Banyo 6.73184 11.82868 Bufonidae Sclerophrys maculata 2009 NMP:P6V 74591/2 0.432 43 Cameroon Ebogo 3.39134 11.46633 Hyperoliidae Hyperolius tuberculatus 2009 NMP:P6V 74561 1.935 44 Cameroon Ebogo 3.39134 11.46633 Phrynobatrachidae Phrynobatrachus auritus 2009 NMP:P6V 74569/1 1715.000 45 Cameroon Ebogo 3.39134 11.46633 Phrynobatrachidae Phrynobatrachus auritus 2009 NMP:P6V 74569/2 157.500 46 Cameroon Ebogo 3.39134 11.46633 Phrynobatrachidae Phrynobatrachus auritus 2009 NMP:P6V 74569/3 61.050 47 Cameroon Ebogo 3.39134 11.46633 Phrynobatrachidae Phrynobatrachus auritus 2009 NMP:P6V 74569/4 2.430 Kedjom Keku 48 Cameroon (= Big Babanki) 6.11620 10.25760 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74612/1 116.500 Kedjom Keku 49 Cameroon (= Big Babanki) 6.11620 10.25760 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74612/2 307.500 Kedjom Keku 50 Cameroon (= Big Babanki) 6.11620 10.25760 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74612/3 6.000 Kedjom Keku 51 Cameroon (= Big Babanki) 6.11620 10.25760 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74612/4 307.000 Kedjom Keku, 52 Cameroon peak 6.09275 10.30315 Arthroleptidae Cardioglossa oreas 2009 NMP:P6V 75162 1.080 53 Cameroon Lake Oku 6.20380 10.45960 Phrynobatrachidae Phrynobatrachus njiomock 2009 NMP:P6V 74523/5 0.944 54 Cameroon Lake Oku 6.20380 10.45960 Phrynobatrachidae Phrynobatrachus njiomock 2009 NMP:P6V 74523/1 1.126 55 Cameroon Mabor 7.70149 12.67672 Bufonidae Sclerophrys maculata 2009 NMP:P6V 74605 81.750 56 Cameroon Mabor 7.70149 12.67672 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74596/3 1.570 57 Cameroon Mabor 7.70149 12.67672 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74586/2 0.759 58 Cameroon Mabor 7.70149 12.67672 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74586/1 1.230 59 Cameroon Mabor 7.70149 12.67672 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74586/4 2.040 60 Cameroon Mabor 7.70149 12.67672 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74586/5 2.200 61 Cameroon Mejung 6.12400 10.24801 Arthroleptidae Leptopelis notatus 2009 NMP:P6V 74608 1370.000 62 Cameroon Mejung 6.12400 10.24801 Ptychadenidae Ptychadena "mascareniensis" D 2009 NMP:P6V 74606/1 337.400 63 Cameroon Mejung 6.12400 10.24801 Ptychadenidae Ptychadena "mascareniensis" D 2009 NMP:P6V 74606/2 5.760

South-West: Bakingili, lava 64 Cameroon flow 4.06810 9.06810 Pipidae Xenopus calcaratus 2016 vgCM16_423 4.883

South-West: Bakingili, lava 65 Cameroon flow 4.06840 9.06820 Pipidae Xenopus allofraseri 2016 vgCM16_406 54.300 South-West: Bakingili, lava 66 Cameroon flow 4.06840 9.06820 Pipidae Xenopus allofraseri 2016 vgCM16_408 12.900

South-West: Bakingili, lava 67 Cameroon flow 4.06840 9.06820 Pipidae Xenopus allofraseri 2016 vgCM16_411 5.105

South-West: Bakingili, lava 68 Cameroon flow 4.06840 9.06820 Pipidae Xenopus allofraseri 2016 vgCM16_412 7.045 69 Cameroon Sud: Mezessé 2.96220 12.15210 Pipidae Xenopus mellotropicalis 2016 vgCM16_389 26.200 70 Cameroon Sud: Mezessé 2.96220 12.15210 Pipidae Xenopus mellotropicalis 2016 vgCM16_390 160.500 71 Cameroon Sud: Mezessé 2.96220 12.15210 Pipidae Xenopus mellotropicalis 2016 vgCM16_392 21.250 72 Cameroon Sud: Onoyong 2.77950 10.82580 Pipidae Xenopus parafraseri 2016 vgCM16_397 90.350 Tchabal 73 Cameroon Gangdaba 7.73035 12.72116 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74576/7 0.822 Tchabal 74 Cameroon Gangdaba 7.73035 12.72116 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74576/8 1.635 Tchabal 75 Cameroon Gangdaba 7.73708 12.72119 Bufonidae Sclerophrys maculata 2009 NMP:P6V 74581 0.831 Tchabal 76 Cameroon Gangdaba 7.74049 12.72305 Arthroleptidae Astylosternus diadematus 2009 NMP:P6V 74589/1 2.090 Tchabal 77 Cameroon Gangdaba 7.74049 12.72305 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74576/1 1.507 Tchabal 78 Cameroon Gangdaba 7.74049 12.72305 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74576/2 1.240 Tchabal 79 Cameroon Gangdaba 7.74049 12.72305 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74576/3 1.032 Tchabal 80 Cameroon Gangdaba 7.74049 12.72305 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74576/4 1.590 Tchabal 81 Cameroon Gangdaba 7.74049 12.72305 Petropedetidae Petropedetes sp. nov. 2009 NMP:P6V 74577/1 3.255 Tchabal 82 Cameroon Gangdaba 7.74049 12.72305 Ranidae Amnirana albolabris 2009 NMP:P6V 74578/1 1.775 Tchabal 83 Cameroon Gangdaba 7.74358 12.71618 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74576/5 0.834 Tchabal 84 Cameroon Gangdaba 7.74358 12.71618 Petropedetidae Petropedetes sp. nov. 2009 NMP:P6V 74577/3 0.821 Tchabal 85 Cameroon Gangdaba 7.74358 12.71618 Petropedetidae Petropedetes sp. nov. 2009 NMP:P6V 74577/2 5.980 Tchabal 86 Cameroon Gangdaba 7.74358 12.71618 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74582/1 0.907 Tchabal 87 Cameroon Gangdaba 7.74358 12.71618 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74582/6 0.326 Tchabal 88 Cameroon Gangdaba 7.74358 12.71618 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74582/2 1.295 Tchabal 89 Cameroon Gangdaba 7.74358 12.71618 Pipidae Xenopus (laevis) poweri 2009 NMP:P6V 74582/3 1.294 Tchabal 90 Cameroon Gangdaba 7.74580 12.71681 Pipidae Xenopus sp. nov. 2009 NMP:P6V 74588/5 0.691 Tchabal 91 Cameroon Gangdaba 7.74580 12.71681 Pipidae Xenopus sp. nov. 2009 NMP:P6V 74588/3 2.130 Tchabal 92 Cameroon Gangdaba 7.74580 12.71681 Pipidae Xenopus sp. nov. 2009 NMP:P6V 74588/6 1.235 Tchabal 93 Cameroon Gangdaba 7.74580 12.71681 Pipidae Xenopus sp. nov. 2009 NMP:P6V 74588/9 2.228 94 Cameroon Touiwa Toumti 7.67093 12.61963 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74574/2 0.967 95 Cameroon Touiwa Toumti 7.67093 12.61963 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74574/1 4.315 96 Cameroon Touiwa Toumti 7.67093 12.61963 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74574/4 4.390 97 Cameroon Touiwa Toumti 7.67093 12.61963 Hyperoliidae Hyperolius riggenbachi 2009 NMP:P6V 74574/5 2.525 98 Cameroon Touiwa Toumti 7.67093 12.61963 Ranidae Amnirana albolabris 2009 NMP:P6V 74575 179.000 Cameroon Gotel Mts., (Nigeria Gangirwal, 99 border) forest 1 7.03071 11.70214 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_021 0.390 Cameroon Gotel Mts., (Nigeria Gangirwal, 100 border) forest 1 7.03071 11.70214 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_018 5.695 Cameroon Gotel Mts., (Nigeria Gangirwal, 101 border) forest 1 7.03071 11.70214 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_019 113.750 Cameroon Gotel Mts., (Nigeria Gangirwal, 102 border) forest 1 7.03071 11.70214 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_025 7.000 Cameroon Gotel Mts., (Nigeria Gangirwal, 103 border) meadow 7.02920 11.69950 Arthroleptidae Leptopelis nordequatorialis 2016 vgCM16_022 2.304 Cameroon Gotel Mts., (Nigeria Gangirwal, site 104 border) 3 7.02920 11.70580 Arthroleptidae Astylosternus rheophilus 2016 vgCM16_023 2.485 Dem. Rep. of 105 the Congo Bazinga 1.4092 28.5723 Hyperoliidae Hyperolius langi 2009 UTEP:22142 38.850 Dem. Rep. of Byonga, Biliki cf. 106 the Congo River -3.34033 28.13085 Hyperoliidae Hyperolius cinnamomeoventris 2010 UTEP:22143 38.930 Dem. Rep. of 107 the Congo Epulu 1.39838 28.56795 Arthroleptidae Leptopelis cf. millsoni 2009 UTEP:22144 45.990 Dem. Rep. of 108 the Congo Nkala -2.58555 16.47182 Arthroleptidae Cardioglossa congolia 2013 UTEP:21111 36.050 Dem. Rep. of road 0.5 km east 109 the Congo of Epulu 1.39693 28.58587 Hyperoliidae Hyperolius bolifambae 2009 UTEP:22145 35.970 MCZ:Herp:A- 110 Gabon Birougou -1.627777778 12.36305556 Arthroleptidae Arthroleptis adelphus 2013 149125 11.604 MCZ:Herp:A- 111 Gabon Birougou -1.627777778 12.36305556 Arthroleptidae Arthroleptis cf. poecilonotus 2013 149156 36.444 MCZ:Herp:A- 112 Gabon Birougou -1.627777778 12.36305556 Arthroleptidae Astylosternus batesi 2013 149158 15.544 MCZ:Herp:A- 113 Gabon Birougou -1.627777778 12.36305556 Arthroleptidae Astylosternus batesi 2013 149160 8.290 MCZ:Herp:A- 114 Gabon Birougou -1.627777778 12.36305556 Arthroleptidae Cardioglossa gracilis 2013 149163 34.108 MCZ:Herp:A- 115 Gabon Birougou -1.627777778 12.36305556 Arthroleptidae Leptopelis ocellatus 2013 149201 23.133 MCZ:Herp:A- 116 Gabon Birougou -1.627777778 12.36305556 Phrynobatrachidae Phrynobatrachus sp. 2013 149215 16.254 MCZ:Herp:A- 117 Gabon Birougou -1.627777778 12.36305556 Ranidae Amnirana amnicola 2013 149167 15.854 MCZ:Herp:A- 118 Gabon Lopé -0.389444444 11.82388889 Arthroleptidae Arthroleptis sylvaticus group 2013 149150 8.002 MCZ:Herp:A- 119 Gabon Lopé -0.36502 11.7906 Phrynobatrachidae Phrynobatrachus auritus 2013 149488 40.739 MCZ:Herp:A- 120 Gabon Lopé -0.36485 11.79053 Phrynobatrachidae Phrynobatrachus auritus 2013 149483 51.647 MCZ:Herp:A- 121 Gabon Lopé -0.36465 11.79062 Phrynobatrachidae Phrynobatrachus auritus 2013 149487 16.677 MCZ:Herp:A- 122 Gabon Lopé -0.36423 11.79073 Phrynobatrachidae Phrynobatrachus auritus 2013 149482 11.927 MCZ:Herp:A- 123 Gabon Lopé -0.36378 11.79065 Phrynobatrachidae Phrynobatrachus auritus 2013 149493 5.396 MCZ:Herp:A- 124 Gabon Lopé -0.36378 11.79068 Phrynobatrachidae Phrynobatrachus auritus 2013 149486 11.993 MCZ:Herp:A- 125 Gabon Lopé -0.3633 11.79027 Phrynobatrachidae Phrynobatrachus auritus 2013 149496 0.162 MCZ:Herp:A- 126 Gabon Lopé -0.3633 11.79027 Phrynobatrachidae Phrynobatrachus auritus 2013 149497 6.396 MCZ:Herp:A- 127 Gabon Lopé -0.36327 11.79018 Phrynobatrachidae Phrynobatrachus auritus 2013 149485 7.409 MCZ:Herp:A- 128 Gabon Lopé -0.36258 11.78953 Phrynobatrachidae Phrynobatrachus auritus 2013 149484 9.659 MCZ:Herp:A- 129 Gabon Minkébé 1.1534 12.69895 Phrynobatrachidae Phrynobatrachus auritus 2013 149414 9.502 MCZ:Herp:A- 130 Gabon Minkébé 1.1536 12.699 Phrynobatrachidae Phrynobatrachus auritus 2013 149415 23.384 MCZ:Herp:A- 131 Gabon Minkébé 1.15383 12.69898 Phrynobatrachidae Phrynobatrachus auritus 2013 149412 34.100 MCZ:Herp:A- 132 Gabon Minkébé 1.15433 12.69897 Phrynobatrachidae Phrynobatrachus auritus 2013 149409 12.927 MCZ:Herp:A- 133 Gabon Minkébé 1.154444444 12.69916667 Arthroleptidae Arthroleptis adelphus 2013 149129 6.351 MCZ:Herp:A- 134 Gabon Minkébé 1.154444444 12.69916667 Arthroleptidae Arthroleptis sylvaticus group 2013 149151 4.640 MCZ:Herp:A- 135 Gabon Minkébé 1.154444444 12.69916667 Arthroleptidae Arthroleptis sylvaticus group 2013 149152 2.767 MCZ:Herp:A- 136 Gabon Minkébé 1.154444444 12.69916667 Arthroleptidae Arthroleptis variabilis 2013 149122 0.557 MCZ:Herp:A- 137 Gabon Minkébé 1.154444444 12.69916667 Arthroleptidae Cardioglossa leucomystax 2013 149164 8.841 MCZ:Herp:A- 138 Gabon Minkébé 1.154444444 12.69916667 Arthroleptidae Leptopelis boulengeri 2013 149198 52.920 MCZ:Herp:A- 139 Gabon Minkébé 1.154444444 12.69916667 Bufonidae Sclerophrys camerunensis 2013 149108 3.425 MCZ:Herp:A- 140 Gabon Minkébé 1.154444444 12.69916667 Hyperoliidae Hyperolius cinnamomeoventris 2013 149189 1.299 141 Gabon Minkébé 1.154444444 12.69916667 Hyperoliidae Hyperolius cinnamomeoventris 2013 MCZ:Cryo:2326 17.196 MCZ:Herp:A- 142 Gabon Minkébé 1.154444444 12.69916667 Hyperoliidae Hyperolius tuberculatus 2013 149191 24.002 MCZ:Herp:A- 143 Gabon Minkébé 1.154444444 12.69916667 Hyperoliidae Hyperolius tuberculatus 2013 149193 13.158 MCZ:Herp:A- 144 Gabon Minkébé 1.154444444 12.69916667 Ptychadenidae Ptychadena aequiplicata 2013 149219 4.729 MCZ:Herp:A- 145 Gabon Minkébé 1.154444444 12.69916667 Ranidae Amnirana amnicola 2013 149172 5.940 MCZ:Herp:A- 146 Gabon Minkébé 1.154444444 12.69916667 Ranidae Amnirana lepus 2013 149184 2.338 MCZ:Herp:A- 147 Gabon Minkébé 1.15495 12.69947 Phrynobatrachidae Phrynobatrachus auritus 2013 149408 18.235 MCZ:Herp:A- 148 Gabon Minkébé 1.155 12.69945 Phrynobatrachidae Phrynobatrachus auritus 2013 149418 25.842 MCZ:Herp:A- 149 Gabon Minkébé 1.15552 12.69752 Phrynobatrachidae Phrynobatrachus auritus 2013 149417 11.952 MCZ:Herp:A- 150 Gabon Ossélé -1.855277778 13.85416667 Arthroleptidae Arthroleptis sylvaticus group 2013 149133 10.629 MCZ:Herp:A- 151 Gabon Ossélé -1.855277778 13.85416667 Arthroleptidae Arthroleptis sylvaticus group 2013 149135 16.480 MCZ:Herp:A- 152 Gabon Ossélé -1.855277778 13.85416667 Arthroleptidae Arthroleptis sylvaticus group 2013 149134 45.269 153 Gabon Ossélé -1.855277778 13.85416667 Arthroleptidae Astylosternus batesi 2013 MCZ:Cryo:2322 18.338 MCZ:Herp:A- 154 Gabon Ossélé -1.855277778 13.85416667 Arthroleptidae Leptopelis notatus 2013 149197 3.392 MCZ:Herp:A- 155 Gabon Ossélé -1.855277778 13.85416667 Bufonidae Sclerophrys gracilipes 2013 149115 6.509 156 Gabon Ossélé -1.855277778 13.85416667 Bufonidae Sclerophrys gracilipes 2013 MCZ:Cryo:2316 8.681 MCZ:Herp:A- 157 Gabon Ossélé -1.855277778 13.85416667 Phrynobatrachidae Phrynobatrachus auritus 2013 149470 25.006 MCZ:Herp:A- 158 Gabon Ossélé -1.855277778 13.85416667 Phrynobatrachidae Phrynobatrachus auritus 2013 149475 28.764 MCZ:Herp:A- 159 Gabon Ossélé -1.855277778 13.85416667 Phrynobatrachidae Phrynobatrachus auritus 2013 149476 11.436 MCZ:Herp:A- 160 Gabon Ossélé -1.855277778 13.85416667 Phrynobatrachidae Phrynobatrachus auritus 2013 149477 19.806 MCZ:Herp:A- 161 Gabon Ossélé -1.855277778 13.85416667 Phrynobatrachidae Phrynobatrachus auritus 2013 149478 9.178 MCZ:Herp:A- 162 Gabon Ossélé -1.855277778 13.85416667 Phrynobatrachidae Phrynobatrachus auritus 2013 149479 19.660 MCZ:Herp:A- 163 Gabon Ossélé -1.855277778 13.85416667 Ptychadenidae Ptychadena "mascareniensis" D 2013 149223 31.895 MCZ:Herp:A- 164 Gabon Ossélé -1.855277778 13.85416667 Ranidae Amnirana albolabris 2013 149175 22.573 MCZ:Herp:A- 165 Gabon Ossélé -1.855277778 13.85416667 Ranidae Amnirana albolabris 2013 149182 24.478 MCZ:Herp:A- 166 Gabon Ossélé -1.855277778 13.85416667 Ranidae Amnirana amnicola 2013 149176 14.280 167 Gabon Ossélé -1.855277778 13.85416667 Ranidae Amnirana sp. 2013 MCZ:Cryo:2325 28.058 MCZ:Herp:A- 168 Gabon Ramba -0.363611111 11.78722222 Arthroleptidae Arthroleptis cf. poecilonotus 2013 149147 25.399 169 Gabon Ramba -0.363611111 11.78722222 Arthroleptidae Leptopelis ocellatus 2013 MCZ:Cryo:2328 18.141 MCZ:Herp:A- 170 Gabon Ramba -0.363611111 11.78722222 Arthroleptidae Scotobleps gabonicus 2013 149225 31.680 171 Gabon Ramba -0.363611111 11.78722222 Arthroleptidae Scotobleps gabonicus 2013 MCZ:Cryo:2334 7.495 MCZ:Herp:A- 172 Gabon Ramba -0.363611111 11.78722222 Ranidae Amnirana albolabris 2013 149168 27.638 MCZ:Herp:A- 173 Gabon Ramba -0.363611111 11.78722222 Ranidae Amnirana albolabris 2013 149169 18.249 Naukluft camp 174 Namibia site -24.26550 16.23950 Pyxicephalidae Amietia poyntoni 2007 AC2969 (lot of 10) N/A Naukluft camp 175 Namibia site -24.26550 16.23950 Pyxicephalidae Amietia poyntoni 2007 AC2969 (lot of 10) N/A Naukluft camp 176 Namibia site -24.26550 16.23950 Pyxicephalidae Amietia poyntoni 2007 AC2969 (lot of 10) N/A Naukluft camp 177 Namibia site -24.26550 16.23950 Pyxicephalidae Amietia poyntoni 2007 AC2969 (lot of 10) N/A Naukluft camp 178 Namibia site -24.26550 16.23950 Pyxicephalidae Amietia poyntoni 2007 AC2969 (lot of 10) N/A 179 Namibia Tsams Ost -24.24689 16.10358 Pyxicephalidae Amietia poyntoni 2007 AC2972 (lot of 10) N/A 180 Namibia Tsams Ost -24.24689 16.10358 Pyxicephalidae Amietia poyntoni 2007 AC2972 (lot of 10) N/A 181 Namibia Tsams Ost -24.24689 16.10358 Pyxicephalidae Amietia poyntoni 2007 AC2972 (lot of 10) N/A 182 Namibia Tsams Ost -24.24689 16.10358 Pyxicephalidae Amietia poyntoni 2007 AC2972 (lot of 10) N/A 183 Namibia Tsams Ost -24.24689 16.10358 Pyxicephalidae Amietia poyntoni 2007 AC2972 (lot of 10) N/A 184 Namibia Tsams Ost -24.24689 16.10358 Pyxicephalidae Amietia poyntoni 2007 AC2972 (lot of 10) N/A 185 Namibia Tsams Ost -24.24689 16.10358 Pyxicephalidae Amietia poyntoni 2007 AC2972 (lot of 10) N/A Rep. of the 186 Congo COPRAGEL -4.589078 12.0556515 Bufonidae Sclerophrys regularis 2010 USNM:584412 0.798 Rep. of the 187 Congo COPRAGEL -4.589078 12.0556515 Hyperoliidae Hyperolius sp. 2010 USNM:584418 0.164 Rep. of the 188 Congo COPRAGEL -4.589078 12.0556515 Hyperoliidae Hyperolius sp. 2010 USNM:584421 0.103 Rep. of the 189 Congo COPRAGEL -4.589078 12.0556515 Hyperoliidae Hyperolius sp. 2010 USNM:584425 0.722 Rep. of the 190 Congo COPRAGEL -4.589078 12.0556515 Hyperoliidae Hyperolius sp. 2010 USNM:584428 0.304 Rep. of the 191 Congo COPRAGEL -4.589078 12.0556515 Hyperoliidae Hyperolius sp. 2010 USNM:584430 0.097 Rep. of the 192 Congo COPRAGEL -4.589078 12.0556515 Hyperoliidae Hyperolius sp. 2010 USNM:584423 1.180 Rep. of the 193 Congo COPRAGEL -4.589078 12.0556515 Hyperoliidae Hyperolius sp. 2010 USNM:584427 1.111 Rep. of the 194 Congo COPRAGEL -4.589078 12.0556515 Ranidae Amnirana sp. 2010 USNM:584438 0.236 Rep. of the 195 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys camerunensis 2008 USNM:576402 42.580 Rep. of the 196 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys camerunensis 2008 USNM:576403 44.490 Rep. of the 197 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys camerunensis 2008 USNM:576404 41.550 Rep. of the 198 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys camerunensis 2008 USNM:576407 32.730 Rep. of the 199 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys camerunensis 2008 USNM:576410 44.450 Rep. of the 200 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys camerunensis 2008 USNM:576412 34.770 Rep. of the 201 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys gracilipes 2008 USNM:576417 40.460 Rep. of the 202 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys gracilipes 2008 USNM:576416 36.580 Rep. of the 203 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys gracilipes 2008 USNM:576242 35.420 Rep. of the 204 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys regularis 2008 USNM:576483 35.900 Rep. of the 205 Congo Impongui 1.0839575 17.300134 Bufonidae Sclerophrys sp. 2008 USNM:herp tissue:4 38.860 Rep. of the 206 Congo Impongui 1.0839575 17.300134 Hyperoliidae Cryptothylax greshoffii 2008 USNM:576609 47.560 Rep. of the 207 Congo Impongui 1.0839575 17.300134 Hyperoliidae Hyperolius dartevellei 2008 USNM:576167 33.420 Rep. of the 208 Congo Impongui 1.0839575 17.300134 Hyperoliidae Hyperolius dartevellei 2008 USNM:576168 32.350 Rep. of the 209 Congo Impongui 1.0839575 17.300134 Hyperoliidae Hyperolius dartevellei 2008 USNM:576180 33.460 Rep. of the 210 Congo Impongui 1.0839575 17.300134 Hyperoliidae Hyperolius sp. 2008 USNM:576183 35.260 Rep. of the 211 Congo Impongui 1.0839575 17.300134 Phrynobatrachidae Phrynobatrachus cf .perpalmatus 2008 USNM:573418 34.920 Rep. of the 212 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576618 36.040 Rep. of the 213 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576629 31.890 Rep. of the 214 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576630 44.270 Rep. of the 215 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576633 35.350 Rep. of the 216 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576638 36.530 Rep. of the 217 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576646 36.970 Rep. of the 218 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576648 33.360 Rep. of the 219 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576649 30.760 Rep. of the 220 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576659 39.770 Rep. of the 221 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576661 34.510 Rep. of the 222 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576662 39.470 Rep. of the 223 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576664 38.480 Rep. of the 224 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576665 40.610 Rep. of the 225 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576674 35.590 Rep. of the 226 Congo Impongui 1.0839575 17.300134 Pipidae Hymenochirus curtipes 2008 USNM:576675 35.220 Rep. of the 227 Congo Impongui 1.0839575 17.300134 Pipidae Xenopus cf. epitropicalis 2008 USNM:573442 36.480 Rep. of the 228 Congo Mayongongo -4.18745 14.95678 Arthroleptidae Arthroleptis sylvaticus 2008 USNM:576272 27.430 Rep. of the 229 Congo Mayongongo -4.18745 14.95678 Bufonidae Sclerophrys maculata 2008 USNM:576449 30.840 Rep. of the 230 Congo Mayongongo -4.18745 14.95678 Bufonidae Sclerophrys regularis 2008 USNM:576566 35.520 Rep. of the 231 Congo Mayongongo -4.18745 14.95678 Bufonidae Sclerophrys sp. 2008 USNM:576392 41.100 Rep. of the 232 Congo Mayongongo -4.18745 14.95678 Hyperoliidae Afrixalus osorioi 2008 USNM:573384 35.660 Rep. of the 233 Congo Mayongongo -4.18745 14.95678 Hyperoliidae Afrixalus sp. 2008 USNM:573401 26.890 Rep. of the 234 Congo Mayongongo -4.18745 14.95678 Hyperoliidae Afrixalus sp. 2008 USNM:573402 30.230 Rep. of the 235 Congo Mayongongo -4.18745 14.95678 Phrynobatrachidae Phrynobatrachus cf .perpalmatus 2008 USNM:576680 28.530 Rep. of the 236 Congo Mayongongo -4.18745 14.95678 Phrynobatrachidae Phrynobatrachus sp. 2008 USNM:576687 31.680 Rep. of the 237 Congo Mayongongo -4.18745 14.95678 Phrynobatrachidae Phrynobatrachus sp. 2008 USNM:576679 30.840 Rep. of the 238 Congo Mayongongo -4.18745 14.95678 Phrynobatrachidae Phrynobatrachus sp. 2008 USNM:576682 21.990 Rep. of the 239 Congo Mayongongo -4.18745 14.95678 Phrynobatrachidae Phrynobatrachus sp. 2008 USNM:576686 44.860 Rep. of the 240 Congo Mayongongo -4.18745 14.95678 Pipidae Xenopus cf. epitropicalis 2008 USNM:573447 31.590 Rep. of the 241 Congo PK-45 -3.917673 15.3952405 Hyperoliidae Afrixalus "quadrivittatus" 2008 USNM:576600 35.970 Rep. of the 242 Congo PK-45 -3.917673 15.3952405 Hyperoliidae Afrixalus "quadrivittatus" 2008 USNM:576602 36.460 Rep. of the Zanaga: 243 Congo Bambama -2.542167 13.5298075 Arthroleptidae Arthroleptis poecilonotus 2010 USNM:584016 0.729 Rep. of the Zanaga: 244 Congo Bambama -2.542167 13.5298075 Arthroleptidae Arthroleptis sylvaticus 2010 USNM:584014 1.143 Rep. of the Zanaga: 245 Congo Bambama -2.542167 13.5298075 Bufonidae Sclerophrys tuberosus 2010 USNM:584136 2.018 Rep. of the Zanaga: 246 Congo Bambama -2.542167 13.5298075 Ranidae Amnirana amnicola 2010 USNM:584215 3.260 Rep. of the USNM:584173 247 Congo Zanaga: Kissiki -2.792862 13.5233355 Phrynobatrachidae Phrynobatrachus batesii 2010 1.062 Rep. of the 248 Congo Zanaga: Kissiki -2.792862 13.5233355 Ranidae Amnirana amnicola 2010 USNM:584213 0.351 Rep. of the 249 Congo Zanaga: Lebayi -2.6768935 13.5871595 Arthroleptidae Arthroleptis poecilonotus 2010 USNM:584055 0.190 Rep. of the 250 Congo Zanaga: Lebayi -2.6768935 13.5871595 Arthroleptidae Astylosternus batesi 2010 USNM:584070 0.685 Rep. of the 251 Congo Zanaga: Lebayi -2.6768935 13.5871595 Arthroleptidae Astylosternus batesi 2010 USNM:584071 0.143 Rep. of the 252 Congo Zanaga: Lebayi -2.6768935 13.5871595 Bufonidae Sclerophrys pusilla 2010 USNM:584150 0.497 Rep. of the 253 Congo Zanaga: Lebayi -2.6768935 13.5871595 Ranidae Amnirana albolabris 2010 USNM:584208 0.260 Rep. of the Zanaga: Congo 254 Simombondo -2.326074 13.6620775 Arthroleptidae Leptopelis aubryioides 2010 USNM:584080 0.786 Rep. of the Zanaga: Congo 255 Simombondo -2.326074 13.6620775 Pipidae Hymenochirus sp. 2010 USNM:584175 0.312 Rep. of the Zanaga: Congo 256 Simombondo -2.326074 13.6620775 Pipidae Xenopus pygmaeus 2010 USNM:584181 0.624 Rep. of the Zanaga: Congo 257 Simombondo -2.326074 13.6620775 Pipidae Xenopus pygmaeus 2010 USNM:584191 0.488 Rep. of the Zanaga: Congo 258 Simombondo -2.326074 13.6620775 Pipidae Xenopus pygmaeus 2010 USNM:584194 0.393 Rep. of the Zanaga: Congo 259 Simombondo -2.326074 13.6620775 Pipidae Xenopus pygmaeus 2010 USNM:584179 1.169 SUPPLEMENTAL MATERIAL S4.— Confirmed positive occurrences for African frogs families with multiple genera. (A) Hyperoliidae; (B) Pyxicephalidae; (C) Arthroleptidae; (D) Pipidae; (E) Bufonidae; (F) Heleophrynidae, and (G) Petropedetidae. Occurrences coded by biogeographical region; map modified from Zimkus et al. 2017.

SUPPLEMENTAL MATERIAL S5.—Aquatic Index of Species Reported with Bd Infection. See Lips et al. (2013) for description of calculation.

Order Family Genus Species Aquatic Index

Anura Alytidae Discoglossus scovazzi 2

Arthroleptidae Arthroleptis adelphus 1

Arthroleptis bioko 1

Arthroleptis perreti 1

Arthroleptis poecilonotus 1

Arthroleptis sylvaticus 1

Arthroleptis tuberosus 1

Arthroleptis variabilis 1

Arthroleptis xenodactyloides 1

Astylosternus batesi 2

Astylosternus diadematus 2

Astylosternus fallax 2

Astylosternus montanus 2

Astylosternus perreti 2

Astylosternus ranoides 2

Astylosternus rheophilus 2

Cardioglossa congolia 2

Cardioglossa elegans 2

Cardioglossa escalerae 2

Cardioglossa gracilis 2

Cardioglossa leucomystax 2

Cardioglossa melanogaster 2

Cardioglossa oreas 2

Leptodactylodon erythrogaster 2

Leptodactylodon mertensi 2.5

Leptopelis aubryi 1.5

Leptopelis argenteus 1.5

Leptopelis aubryioides 1.5

Leptopelis bocagii 1.5

Leptopelis boulengeri 1.5

Leptopelis brevirostris 1.5 Leptopelis calcaratus 1.5

Leptopelis christyi 1.5

Leptopelis gramineus 1.5

Leptopelis karissimbensis 1.5

Leptopelis kivuensis 1.5

Leptopelis millsoni 1.5

Leptopelis modestus 1.5

Leptopelis mtoewaate 1.5

Leptopelis nordequatorialis 1.5

Leptopelis notatus 1.5

Leptopelis occelatus 1.5

Leptopelis palmatus 1.5

Leptopelis parbocagii 1.5

Leptopelis ragazzii 1.5

Leptopelis rufus 1.5

Leptopelis spiritusnoctis 1.5

Leptopelis vannutellii 1.5

Leptopelis viridis 1.5

Leptopelis xenodactylus 1.5

Nyctibates corrugatus 2

Scotobleps gabonicus 2

Trichobatrachus robustus 2

Breviceptidae Balebreviceps hillmani 1

Bufonidae Altiphrynoides malcolmi 1

Nectophryne afra 1.5

Nectophryne batesii 1.5

Nectophrynoides asperginis 1

Sclerophrys camerunensis 1.5

Sclerophrys funerea 1.5

Sclerophrys gracilipes 1.5

Sclerophrys gutturalis 1.5

Sclerophrys maculata 1.5

Sclerophrys poweri 1.5

Sclerophrys pusilla 1.5 Sclerophrys regularis 1.5

Sclerophrys tuberosus 1.5

Vandijkophrynus gariepensis 1.5

Vandijkophrynus robinsoni 1.5

Wolsterstorffina mirei 1.5

Wolterstorffina parvipalmata 1.5

Conrauidae Conraua crassipes 3

Dicroglossidae Hoplobatrachus occipitalis 2

Heleophrynidae Hadromophryne natalensis 2.5

Heleophryne hewitti 2.5

Heleophryne orientalis 2.5

Heleophryne purcelli 2.5

Heleophryne regis 2.5

Heleophryne rosei 2.5

Hylidae Hyla meridionalis 1.5

Hyperoliidae Afrixalus clarkei 1.5

Afrixalus crotalus 1.5

Afrixalus dorsalis 1.5

Afrixalus enseticola 1.5

Afrixalus fornasini 1.5

Afrixalus fulvovittatus 1.5

Afrixalus lacteus 1.5

Afrixalus laevis 1.5

Afrixalus nigeriensis 1.5

Afrixalus osorioi 1.5

Afrixalus paradorsalis 1.5

Afrixalus quadrivittatus 1.5

Afrixalus spinifrons 1.5

Afrixalus sylvaticus 1.5

Afrixalus uluguruensis 1.5

Afrixalus wittei 1.5

Alexteroon hypsiphonus 1.5

Alexteroon obstetricans 1.5

Callixalus pictus 1.5 Chrysobatrachus cupreonitens 1.5

Cryptothylax greshoffii 1.5

Hyperolius acuticeps 2

Hyperolius adametzi 2

Hyperolius adspersus 2

Hyperolius balfouri 2

Hyperolius bolifambae 2

Hyperolius camerunensis 2

Hyperolius castaneus 2

Hyperolius cinnamomeoventris 2

Hyperolius concolor 2

Hyperolius constellatus 2

Hyperolius dartevellei 2

Hyperolius discodactylus 2

Hyperolius endjami 2

Hyperolius frontalis 2

Hyperolius fusciventris 2

Hyperolius glandicolor 2

Hyperolius hutsebauti 2

Hyperolius kivuensis 2

Hyperolius koehleri 2

Hyperolius langi 2

Hyperolius lateralis 2

Hyperolius molleri 2

Hyperolius nasicus 2

Hyperolius nasutus 2

Hyperolius ocellatus 2

Hyperolius pardalis 2

Hyperolius phantasticus 2

Hyperolius pickersgilli 2

Hyperolius picturatus 2

Hyperolius pictus 2

Hyperolius pseudargus 2

Hyperolius puncticulatus 2 Hyperolius pusillus 2

Hyperolius quinquevittatus 2

Hyperolius riggenbachi 2

Hyperolius robustus 2

Hyperolius spinigularis 2

Hyperolius sylvaticus 2

Hyperolius thomensis 2

Hyperolius tuberculatus 2

Hyperolius tuberilinguis 2

Hyperolius viridiflavus 2

Hyperolius viridistriatus 2

Kassina decorata 2

Kassina maculosa 2

Kassina senegalensis 2

Opisthothylax immaculatus 1.5

Paracassina obscura 1.5

Phlyctimantis leonardi 1.5

Phlyctimantis verrucosus 1.5

Microhylidae Phrynomantis bifasciatus 1.5

Pelobatidae Pelobates varavaldii 1.5

Petropedetidae Arthroleptides yakusini 2

Ericabatrachus baleensis 1.5

Petropedetes euskircheni 1.5

Petropedetes newtoni 2

Petropedetes palmipes 3

Phrynobatrachidae Phrynobatrachus acridoides 2

Phrynobatrachus acutirostris 2

Phrynobatrachus africanus 2

Phrynobatrachus asper 2

Phrynobatrachus auritus 2

Phrynobatrachus calcaratus 2

Phrynobatrachus cornutus 2

Phrynobatrachus cricogaster 2

Phrynobatrachus dendrobates 2 Phrynobatrachus dispar 2

Phrynobatrachus graueri 2

Phrynobatrachus jimzimkusi 2

Phrynobatrachus krefftii 2

Phrynobatrachus leveleve 2

Phrynobatrachus liberiensis 2

Phrynobatrachus minutus 2

Phrynobatrachus natalensis 2

Phrynobatrachus natalensis 2

Phrynobatrachus njiomock 2

Phrynobatrachus plicatus 2

Phrynobatrachus steindachneri 2

Phrynobatrachus versicolor 2

Phrynobatrachus werneri 2

Pipidae Hymenochirus curtipes 3

Xenopus allofraseri 3

Xenopus andrei 3

Xenopus borealis 3

Xenopus calcaratus 3

Xenopus clivii 3

Xenopus epitropicalis 3

Xenopus eysoole 3

Xenopus sp. (amieti group) 3

Xenopus gilli 3

Xenopus laevis 3

Xenopus longipes 3

Xenopus mellotropicalis 3

Xenopus muelleri 3

Xenopus parafraseri 3

Xenopus poweri 3

Xenopus pygmaeus 3

Xenopus tropicalis 3

Xenopus victorianus 3

Xenopus wittei 3 Ptychadenidae Ptychadena aequiplicata 2

Ptychadena anchietae 2

Ptychadena bibroni 2

Ptychadena chrysogaster 2

Ptychadena erlangeri 2

Ptychadena longirostris 2 Ptychadena mascareniensis 2 (complex)

Ptychadena neumanni 2

Ptychadena newtoni 2

Ptychadena oxyrhynchus 2

Ptychadena perreti 2

Ptychadena pumilio 2

Ptychadena uzungwensis 2

Pyxicephalidae Amietia angolensis 2

Amietia delalandii 2

Amietia fuscigula 2

Amietia hymenopus 2

Amietia johnstoni 2

Amietia poyntoni 2

Amietia ruwenzorica 2

Amietia vertebralis 3

Amietia wittei 2

Cacosternum aggestum 2

Cacosternum boettgeri 2

Cacosternum striatum 2

Microbatrachella capensis 2

Nothophryne broadleyi 1.5

Strongylopus fasciatus 1.5

Strongylopus fuelleborni 1.5

Strongylopus grayii 1.5

Tomopterna cryptotis 1.5

Tomopterna natalensis 1.5

Ranidae Amnirana albolabris 2 Amnirana amnicola 2

Amnirana lepus 2

Amnirana longipes 2

Rhacophoridae Chiromantis rufescens 1.5

Gymnophiona Dermophiidae Geotrypetes seraphini 1

Schistometopum thomense 1

Herpelidae Boulengerula uluguruensis 1

Herpele squalostoma 1

Indotyphlidae Idiocranium russeli 1

Scolecomorphidae Crotaphatrema lamottei 1

Scolecomorphus kirkii 1

SUPPLEMENTAL MATERIAL S6.—Graph of confirmed positive occurrences for African caecilian families with multiple genera. (A) Scolecomorphidae; (B) Herpelidae; and (C) Dermophiidae. Occurrences coded by biogeographical region; map modified from Zimkus et al. 2017.

SUPPLEMENTAL MATERIAL S7.—Details of the contributions of each variable (A-Q) to the calculated ENMs, showing the mean individual response curves (red) and their standard deviation (blue).

SUPPLEMENTAL MATERIAL S8.—Estimated relative variable contribution without (percentage contribution) and with random permutation (permutation importance) of the values. Variable Percent Permutation contribution importance

Rain low 30.6 1.2

Elev. var. 23.9 10.1

MODIS tree 9.4 4.1

Temp. max low 8.0 31.3

MODIS bare 5.3 2.2

Temp. max high 4.8 2.7

Elev. contr. 4.7 0.5

SPOT 4 3 3.5 7.7

Temp. min high 2.6 1.9

Rain total 2.3 2.7

MODIS herb 1.6 17.2

Temp. min SD 0.9 5.8

SPOT 4 2 0.6 0.6

Temp. max SD 0.6 9.3

Temp. min low 0.4 0.2

Rain SD 0.4 0.9

Rain max 0.3 1.6

SUPPLEMENTAL MATERIAL S9.—Test omission rate and predicted area as a function of the cumulative threshold, averaged over the replicate runs.

SUPPLEMENTAL MATERIAL S10.—Receiver operating characteristic (ROC) curve averaged over the replicate runs.

SUPPLEMENTAL MATERIAL S11.—Results of the jackknifing tests of variable importance for training data.

SUPPLEMENTAL MATERIAL S12.—Results of the jackknifing tests of variable importance for testing data.

SUPPLEMENTAL MATERIAL S13.—Results of the jackknifing of AUC for Bd.