Predicting the Distribution of the Amphibian Pathogen Batrachochytrium dendrobatidis in the New World1

One application of ecological niche modeling is predicting suitable areas for the establishment of invasive species. Herein, I model the fundamental niche of the chytrid fungus Batrachochytrium dendrobatidis, a pathogen linked to amphibian declines on several continents. Niche models were generated with the Genetic Algorithm of Rule‐Set Prediction using point distribution data of the pathogen and digital maps of environmental variables integrated in a GIS environment. The distribution of regions suitable for B. dendrobatidis in the New World is extensive and includes significant portions of: (1) Sierra Madre Occidental pine‐oak forest; (2) Sonoran and Sinaloan dry forest; (3) Veracruz moist forest; (4) Central America east from the Isthmus of Tehuantepec; (5) Caribbean Islands; (6) temperate forest in Chile and western Argentina south of latitude 30°S; (7) Andes above 1000 m of altitude in Venezuela, Colombia, and Ecuador; (8) eastern slopes of the Andes in Peru and Bolivia; (9) Brazilian Atlantic forest; (10) Uruguay, Paraguay, and northeastern Argentina; (11) southwestern and Madeira‐Tapajós Amazonian tropical rainforests. The regions with the highest suitability for B. dendrobatidis include habitats that contain the world's most diverse amphibian faunas. Models were built with New World localities, but also showed strong predictability for B. dendrobatidis localities in the Old World. Out of a total of 59 reported Old World localities for B. dendrobatidis, 56 occurred within regions with high predicted suitability. I also present analyses of the environmental envelope of B. dendrobatidis and discuss the implications of the results for the conservation of amphibians in the neotropics.     

Cross‐scale effects of structural and functional connectivity in pond networks on amphibian distribution in agricultural landscapes

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Biogeography of wild <Emphasis Type="Italic">Arachis</Emphasis> (<Emphasis Type="Italic">Leguminosae</Emphasis>):distribution and environmental characterisation

Geographic Information System (GIS) tools are applied to a comprehensive database of 3514 records of wild Arachis species to assist in the conservation and utilisation of the species by: (a) determining the distributional range of species and their abundance; (b) characterising species environments; (c) determining the geographical distribution of species richness; and (d) determining the extent to which species are associated with river basins. Distributional ranges, climatic variables and indices of endemism for each species are tabulated. A. duranensis Krapov. & W.C. Gregory, the most probable donor of the A genome to the cultivated peanut, is distributed in close proximity to both the proposed donor of the B genome, A. ipaënsis, and the closest wild relative of the cultigen, A. monticola Krapov. & Rigoni. This region in the eastern foothills of the Andes and the adjoining chaco regions of Argentina, Bolivia and Paraguay, is a key area for further exploration for wild Arachis. An area of particularly high species richness occurs in the State of Mato Grosso, close to the Gran Pantanal in southwest Brazil. Seventy-one percent of the species were found to have some degree of association with water catchment areas, although in most cases it was difficult to determine whether this was due to climatic adaptation reasons, restricted dispersal due to geocarpic habit, or the role of watercourses as a principal dispersal agent. In only two cases could climatic adaptation be eliminated as the reason for species distribution.     

Chytridiomycosis Survey in Wild and Captive Mexican Amphibians

Mexico, a rich country in terms of amphibian diversity, hosts about 375 described species. Population declines have been documented for several species where it is evident that their habitat is being destroyed or modified. However, other species which inhabit pristine areas are declining as well. It has been suggested that the chytrid fungus Batrachochytrium dendrobatidis (B.d.) may be one of the causes of the enigmatic declines in Mexico. We surveyed a total of 45 localities, in 12 states across Mexico, examining a total of 360 specimens representing 14 genera and 30 species. We also examined 91 specimens of Ambystoma mexicanum from a captive population in Mexico City as well as one Pachymedusa dacnicolor obtained in a pet shop. We used a two-tiered technique to detect the pathogen. For wild-caught specimens, we utilized light microscopy to identify presence of B.d. sporangia in amphibian skin. Then, to verify the infection, we used a quantitative real-time PCR assay on collected skin sections which is specific for B.d. For captive animals, we used a nonlethal version of the real-time PCR technique. We found evidence of B.d. infection in 111 animals comprising 14 species in 13 localities. A large percentage (84%) of Ambystoma mexicanum from the colony were infected with B.d. The two most highly infected individuals were the endangered Ambystoma mexicanum, from a captive colony, and Pachymedusa dacnicolor, purchased at a pet shop. The online version of this article (doi:) contains electronic supplementary material, which is available to authorized users.     

Distribution models for the amphibian chytrid Batrachochytrium dendrobatidis in Costa Rica: proposing climatic refuges as a conservation tool

Aim We use novel data on the occurrence of the amphibian chytrid fungus Batrachochytrium dendrobatidis in Costa Rica to model its potential distribution in that country. Location Lowland and montane areas of Costa Rica. Methods We use published and new data on the presence of B. dendrobatidis on 647 amphibians (35 species). Screening was performed through histological techniques by which 156 sites were surveyed. Of these, 21 were found to have the amphibian chytrid. Maxent, a presence‐only distribution modelling technique, was used to create 100 predictions of B. dendrobatidis occurrence, of which the most accurate 10 (based on area under the receiver‐operating characteristic curve) were chosen to create a composite distribution model. This approach increased confidence in model predictions, distinguishing areas of high probability of occurrence and low variability across model runs (higher confidence) from those with high probability but high variability (lower confidence). Results Predicted distribution patterns were not uniform along Costa Rica's mountains, where most amphibian declines have occurred. The pathogen was predicted to occur with greater probability on the Caribbean slopes than on the Pacific slopes. While high temperature seems to constrain the distribution of the pathogen, areas that also have small amounts of rainfall during the driest period of the year were predicted to have low probability of B. dendrobatidis occurrence. Main conclusions The model predicts that the Santa Elena Peninsula and the Central Valley have low probabilities of B. dendrobatidis occurrence, suggesting that they could function as refuges for amphibians. In such refugial areas, one could expect B. dendrobatidis to be absent, or to be present in low abundance (rendering an epidemic outbreak of chytridiomycosis unlikely). Craugastor ranoides, which belongs to a group of frogs particularly sensitive to chytridiomycosis outbreaks, persists in the hot and seasonally dry Santa Elena Peninsula but disappeared in the nearby colder and more humid Guanacaste Volcanic Chain. This information suggests that climatic refuges, where environmental conditions prevent disease outbreaks, could be an important component in amphibian conservation.     

Host range and geographical distribution of <Emphasis Type="Italic">Ixodes sigelos</Emphasis> (Acari: Ixodidae)

Larvae, nymphs and females of Ixodes sigelos Keirans, Clifford and Corwin (Ixodidae) were collected in 13 localities of the Patagonian region of Argentina parasitizing eight species of sigmodontine rodents. We report for the first time adults of I. sigelos for Argentina. Besides, we extend the southern limit of its geographical distribution, and six species of sigmodontines are added as new host species of I. sigelos (Phyllotis xanthopygus, Euneomys chinchilloides, Calomys musculinus, Reithrodon auritus, Loxodontomys micropus and Eligmodontia morgani). The presence of larvae, nymphs and females on sigmodontines, as well as more than 50% of the individuals engorged, indicate that I. sigelos can develop the complete parasitic phase of its life cycle on these small mammals. The geographical distribution of I. sigelos is restricted to the biogeographical Andean Region in Argentina and Chile.     

The genus Bipinnula (Orchidaceae: Chloraeinae) in Argentina

A taxonomic synopsis of Bipinnula Comm. ex Juss. (Orchidaceae: Chloraeinae) in Argentina is presented. Three species are recognized: B. biplumata Rchb.f., B. penicillata (Rchb.f.) Cisternas & Salazar and B. polysyka Kraenzl. A key to separate these species is presented and their distribution in Argentina is presented. All species are described in detail and photographs of diagnostic vegetative and floral features, as well as distributional, ecological and nomenclatural comments are included. Lectotypes for the basionyms Arethusa biplumata L.f., Chloraea arechavaletae Kraenzl. and B. polysyka Kraenzl. are designated. All studied species are terrestrial orchids that grow in grasslands of the heavily transformed Pampas biome; two of them are restricted to a few counties. Therefore, an assessment on their conservation status is urgently needed.     

First Evidence of <Emphasis Type="Italic">Batrachochytrium dendrobatidis</Emphasis> in China: Discovery of Chytridiomycosis in Introduced American Bullfrogs and Native Amphibians in the Yunnan Province,China

Although the chytrid fungus Batrachochytrium dendrobatidis (Bd), the etiological agent of amphibian chytridiomycosis, has been implicated in mass mortality and population declines on several continents around the world, there have been no reports on the presence of Bd infections in amphibians in China. We employed quantitative PCR and histological techniques to investigate the presence of Bd in introduced North American bullfrogs (Rana catesbeiana) (referred to hereafter as bullfrog) and native amphibians in bullfrog-invaded areas of the Yunnan Province, China. A total of 259 samples at five wild sites were collected between June and September in 2007 and 2008, including bullfrogs and four native amphibian species (Rana pleuraden, Rana chaochiaoensis, Odorrana andersonii, and Bombina maxima). In addition, 37 samples of adult bullfrogs were obtained from a food market. Bd infections were discovered in bullfrogs and three native amphibian species from all of the surveyed sites. Of the 39 Bd-positive samples, 35 were from wild-caught bullfrog tadpoles, postmetamorphic bullfrogs, R. pleuraden, R. chaochiaoensis, and O. andersonii, and four were from adult bullfrogs from the market. Our results provide the first evidence of the presence of Bd in Chinese amphibians, suggesting that native amphibian diversity in China is at risk from Bd. There is an urgent need to monitor the distribution of Bd in amphibians in China and understand the susceptibility of native amphibian species to chytridiomycosis. Strict regulations on the transportation of bullfrogs and the breeding of bullfrogs in markets and farms should be drafted in order to stop the spread of Bd by bullfrogs.     

First record of the avian ectoparasite <Emphasis Type="Italic">Philornis downsi</Emphasis> Dodge &; Aitken, 1968 (Diptera: Muscidae) in Argentina

Species of Philornis Meinert, 1890 (Diptera, Muscidae) are Neotropical dipterans that include species with parasitic larvae which feed on nestling birds. To date, all Philornis species that have been recorded from Argentina have parasitic subcutaneous larvae. Here, for the first time for Argentina, we report the finding of Philornis downsi Dodge & Aitken, 1968, a fly with a nest-dwelling, semi-haematophagous larva. This record, from the humid Chaco ecoregion of Argentina in the nest of a saffron finch Sicalis flaveola pelzelni Sclater, substantially extends the known distribution of this species. We also report the consensus sequences of the internal transcribed spacer 1 (ITS1) and ITS2 regions of three of the specimens for future reference and comparison. Further investigation is needed to determine whether Argentina is part of the historical range of P. downsi or, alternatively, represents a recent expansion of its range, perhaps due to climatic changes or other factors of global environmental variation.     

Establishing species' environmental requirements to understand how the southernmost species of South American pitvipers (Bothrops,Viperidae) are distributed: A niche‐based modelling approach

Three Bothrops species are known to be present along an extensive and generally xeric band of the south‐eastern portion of South America. Yet, the environmental factors responsible for the maintenance of the structure of this community have remained undetermined. To have a better understanding of snakes' geographic range limits in Argentina, we modelled the ecological niche of Bothrops alternatus, B. ammodytoides and B. diporus to identify areas of their occurrence and sympatry. We used snakes presence records and pseudo‐absence data with topographic (altitude and slope), vegetation and 19 bioclimatic variables as predictors. These variables were evaluated for each species, using Multiple Logistic Regression Analysis. The distribution of B. alternatus was correlated with temperature, precipitation and vegetation; that of B. ammodytoides with altitude, slope, precipitation, temperature and vegetation; and that of B. diporus related only to precipitation and temperature variables. Five variables appeared in all three models (i.e. annual precipitation, precipitation seasonality, precipitation of the driest month, mean diurnal temperature range and temperature range), although with different effects in different species. Probable sympatry areas were identified, for B. alternatus – B. ammodytoides in a narrow area, for B. diporus – B. alternatus in a wide area and B. ammodytoides – B. diporus in a small area. While some factors constrain the distribution of the southernmost pitvipers community in South America to spatially non‐overlapping ranges, others facilitate sympatry by allowing spatial coexistence. We found high sympatry between phylogenetically distantly related pitvipers. Niche‐based Geographic Information Systems modelling allowed us to identify the environmental factors correlated with each pitviper range in Argentina and in its contact zones, and to understand species distributional limits.     

<Emphasis Type="Italic">Coccocarpia tomentosa</Emphasis>, una nueva especie de Argentina (Ascomycetes liquenizados,Peltigerales, Coccocarpiaceae)

In this paper Coccocarpia tomentosa, from Misiones, Argentina, is described and illustrated. Comments on five related species also known from Paraguay and Brazil are included. The known distribution of C. pellita is extended and it is cited for the first time from Argentina.     

Amphibian assemblages and diversity patterns in two forest ecosystems of South-Eastern Nigeria

Despite recent anthropogenic interference, south-eastern (SE) Nigeria historically hosts a myriad of pristine forest habitats that could sustain diverse amphibian assemblages. Currently, land conversion of primary forests through human activities has led to modified habitats, such as agricultural fields and secondary forest. This calls for biodiversity documentation and initiation of conservation plans. However, the dearth of herpetological studies has limited knowledge of the amphibian species richness and distribution patterns in some forest ecosystems of SE Nigeria. Thus, we surveyed two less-explored intact forests in SE Nigeria—Afi Forest Reserve (AFR) and Ikpan forest ecosystem (IFE)—to document the patterns of amphibian diversity and distribution. Specimens were searched for opportunistically using visual and acoustic survey methods. Literature searches were conducted to document additional species that were not encountered. Using the observational records and arrays of diversity indices, the patterns of species richness and abundance across habitat types were evaluated. The result of our study revealed the presence of 30 and 31 amphibian species from AFR and IFE, respectively. Our field collections included two unidentified species (Sclerophrys sp. and Hyperolius sp.) from AFR and one (Phrynobatrachus sp.) from IFE. Analyses revealed high species diversity and richness in forest and swamps compared with the agricultural field and savannah. Given the increased human activities and the antecedent threats to forest ecosystems of SE Nigeria, we offer recommendations for the initiation of conservation plans for these two surveyed localities.     

Phylogeographic analysis reveals northerly refugia for the riverine amphibian Triturus dobrogicus (Caudata: Salamandridae)

We investigated the recent evolutionary history of the Danube crested newt, Triturus dobrogicus through reconstructions of: (1) the number and position of refugia at the last glacial maximum, (2) the role of major central European rivers in pattern of post‐glacial dispersal, and (3) the present‐day distribution pattern. We analysed sequences of mitochondrial DNA (ND2, 1065 bp) and six microsatellite loci in 363 T. dobrogicus individuals from 58 populations covering the range of the species. Our analyses suggested that T. dobrogicus survived the last glacial maximum in two separate refugia positioned in northwestern Pannonia and in Southern Pannonia from where its range expanded along the Danube and Tisza Rivers. Our findings also confirmed that rivers played an important role in shaping the evolutionary history of amphibian species in Central Europe. We compared the T. dobrogicus range with another lowland amphibian, the fire‐bellied toad Bombina bombina, using species distribution modelling. In line with these models, the isolated mountains inside Pannonia are occupied not by B. bombina, but by its mountain‐dwelling sister‐species B. variegata. However, in contrast to the model, crested newts (including T. dobrogicus) are absent from these mountains. We attribute this biogeographical discrepancy to the positioning of the species’ refugia at the last glacial maximum.     

Diversity of form in the amphibian papilla of Puerto Rican frogs

Summary In modern frogs, the amphibian papilla exhibits a caudal extension whose shape, relative length, and proportion of hair cells vary markedly from species to species. Tuning in the caudal extension is organized tonotopically and evidently involves the tectorium. In terms of the proportion of amphibian-papillar hair cells in the caudal extension, we report more diversity among 8 species of a single genus (Eleutherodactylus) on a single island (Puerto Rico) than has been found so far among all of the (more than 50) other modern anurans examined for this feature from around the world. These 8 Puerto Rican species have overlapping habitat and conspicuous diversity in the male advertisement call. For 7 of the 8 species, we report that the call has transient spectral components in the frequency range of the amphibian papilla, and that the proportion of caudal extension hair cells and the frequency distribution of those components are correlated. Thus one might conclude that the selective pressures that led to diversity of calls among the 8 species also led to diversity in form of the amphibian papilla.Abbreviations AP amphibian papilla - BEF best excitatory frequency - PR Puerto Rican - SEM scanning electron microscope - SVL snout-vent length - TC tectorial corner - TM tectorial membrane     

Three new species of Euathlus Ausserer, 1875 (Araneae,Theraphosidae) from Argentina

ABSTRACT

Euathlus

Ausserer, 1875 is a South American genus of spiders of the family Theraphosidae known from Chile and Argentina. Three new species from Argentina: Euathlus mauryi sp. n. (from San Juan province), Euathlus grismadoi sp. n. (from La Rioja province), and Euathlus pampa sp. n. (from Salta province) are described here. New records of the previous species Euathlus diamante and Euathlus tenebrarum are contributed. The distribution of these new species expands the geographical distribution of the genus along the Andean hills, with Salta province as the northernmost record of the genus. A key is provided for identification of Euathlus species as well as a map with all the records in Argentina.     

Heavy metal pollution negatively correlates with anuran species richness and distribution in south‐eastern Australia

Heavy metal pollution has likely played an important role in global biodiversity decline, but there remains a paucity of information concerning the effects of metals on amphibian diversity. This study assessed anuran species richness and distribution in relation to sediment metal content and water chemistry in wetlands located along the Merri Creek corridor in Victoria, south‐eastern Australia. Anurans were present in 60% (21/35) of study sites, with a total of six species detected: the eastern common froglet (Crinia signifera), the eastern sign‐bearing froglet (Crinia parinsignifera), the southern brown tree frog (Litoria ewingii), the growling grass frog (Litoria raniformis), the eastern banjo frog (Limnodynastes dumerilii) and the spotted marsh frog (Limnodynastes tasmaniensis). Mean species richness was 1.77 ± 0.32 per site, and species richness ranged from zero to six species per site. Across sites, species richness correlated negatively with sediment concentrations of six heavy metals: copper, nickel, lead, zinc, cadmium and mercury. Species richness also correlated negatively with wetland water electrical conductivity (a proxy for salinity) and concentrations of orthophosphate. Distributions of the three most commonly observed frog species (C. signifera, L. tasmaniensis and L. ewingii) were significantly negatively associated with the total level of metal contamination at individual sites. The study is the first to provide evidence for an association between metal contamination and anuran species richness and distribution in the southern hemisphere, adding to a small but growing body of evidence that heavy metal pollution has contributed to global amphibian decline.     

Amphibian Pathogen Batrachochytrium dendrobatidis Is Inhibited by the Cutaneous Bacteria of Amphibian Species

Population declines of amphibian species in many parts of the world are associated with a lethal fungal pathogen, Batrachochytrium dendrobatidis. Using laboratory challenge assays, we describe the inhibition of B. dendrobatidis by members of eight genera of bacteria isolated from the skin of two amphibian species that exhibit parental care behavior (Plethodon cinereus and Hemidactylium scutatum). We found that members of three genera of bacteria isolated from the skins of the salamander P. cinereus and members of seven genera isolated from the salamander H. scutatum inhibited the growth of B. dendrobatidis. Understanding how B. dendrobatidis interacts with an ecological community of cutaneous flora may be important in explaining and preventing amphibian population declines.     

Experimental Infection and Repeat Survey Data Indicate the Amphibian Chytrid Batrachochytrium dendrobatidis May Not Occur on Freshwater Crustaceans in Northern Queensland, Australia

Chytridiomycosis is a fatal disease of amphibians, caused by the amphibian chytrid Batrachochytrium dendrobatidis. The disease is unusual in that it may drive many amphibian species to local extinction during outbreaks. These dramatic declines in host population numbers could be facilitated if the pathogen can grow as a saprobe or on alternative hosts, a feature common to other chytrid species. This is also supported by in vitro work that demonstrates B. dendrobatidis can grow and reproduce in the absence of amphibian cells. In a previous study, B. dendrobatidis was detected on freshwater shrimp from rain forest streams in northern Queensland, Australia, using diagnostic PCR. We set out to confirm and further investigate the presence of B. dendrobatidis on crustaceans by carrying out more extensive sampling of shrimp in the field, experimental B. dendrobatidis infection trials using shrimp and crayfish, and PCR verification of the presence of B. dendrobatidis from shrimp samples that previously tested positive. We could not confirm the presence of B. dendrobatidis on shrimp, and report that original positive tests in shrimp reported by Rowley et al. (2006) were likely false. Thus, we suggest that shrimp may not be an important reservoir host for B. dendrobatidis.     

Conservation and historical distribution of two bumblebee species from the Atlantic Forest

In this study, we analysed the processes resulting in the origin of two endemic sister species of bumblebees in the Brazilian Atlantic Forest. We studied the historical distribution pattern of Bombus bahiensis, which is restricted to small fragments in eastern Brazil and the phylogeographic pattern and historical demography of B. brasiliensis, which is widely distributed in southern and south-eastern Brazil and neighbouring regions of Uruguay, Paraguay, and Argentina. We used ecological niche models, niche analyses, and genetic and distribution data (i) to test the role of niche differentiation on the divergence between the two species, (ii) to find potential distribution areas for the most restricted B. bahiensis, and (iii) to evaluate the conservation status of both species. Our results showed that B. brasiliensis populations are able to disperse across mosaics of anthropogenic and preserved areas and exhibit low levels of spatial genetic structure. Otherwise, B. bahiensis presented a restricted distribution range and likely a lower diversity, where it is suffering with an increasing habitat loss. The climatic oscillations of the Pleistocene influenced the population structure of both species in different ways, probably due to differences in their effective population sizes, physiology and past demographic fluctuations. Specifically, while B. brasiliensis expanded its distribution range in the last 500,?000?years throughout most of the Atlantic Forest, B. bahiensis remained restricted to a small rainforest area between southern Bahia and northern Espírito Santo states in Brazil. In its southern distribution, in the state of Espírito Santo, B. bahiensis is currently very rare or extinct. Currently, the adjacent ranges of Bombus brasiliensis and B. bahiensis do not overlap and our results indicate that these species may have further diversified through a reinforcement process associated to niche specialization and differentiation.     

Botryocladius gen.n.: a new transantarctic genus of orthocladiine midge (Diptera: Chironomidae)

A new genus of orthocladiine Chironomidae, Botryocladius (type species B. grapeth sp.n. from eastern Australia) is described and illustrated in all life history stages. All thirteen included species are described as new, six from eastern Australia (B. grapeth, B. brindabella, B. mdfrc, B. collessi, B. tasmania, B. australoalpinus), two from Western Australia (B. bibulmun and B. freemani), one from ephemeral streams in Australia (B. petrophilus) and four from Patagonian Argentina and Chile (B. edwardsi, B. glacialis, B. mapuche and B. tronador). All Australian species are known from at least pupal exuviae, most from adult males and several from larvae. In contrast, only B. edwardsi amongst Neotropical species is known from the adult male; all others are described from pupal exuviae. The immature stages are lotic in Australian permanent and temporary streams and Patagonian glacial streams and rivers, and lentic in Neotropical glacial-fed and Australian subalpine lakes. Botryocladius appears to belong with a grouping centred on two formally undescribed taxa from Australia. The genus evidently demonstrates a vicariant distribution with at least two sister-group relationships between South American and Australian taxa, providing a minimum dating for the clade of 38 Ma., with apparent absence from New Zealand indicating a maximum date of 80 Ma.