Steffen Reichle

Systematics of spiny-backed treefrogs (Hylidae: Osteocephalus): An Amazonian puzzle

Spiny‐backed tree frogs of the genus Osteocephalus are conspicuous components of the tropical wet forests of the Amazon and the Guiana Shield. Here, we revise the phylogenetic relationships of Osteocephalus and its sister group Tepuihyla, using up to 6134 bp of DNA sequences of nine mitochondrial and one nuclear gene for 338 specimens from eight countries and 218 localities, representing 89% of the 28 currently recognized nominal species. Our phylogenetic analyses reveal (i) the paraphyly of Osteocephalus with respect to Tepuihyla, (ii) the placement of ‘Hyla’ warreni as sister to Tepuihyla, (iii) the non‐monophyly of several currently recognized species within Osteocephalus and (iv) the presence of low (<1%) and overlapping genetic distances among phenotypically well‐characterized nominal species (e.g. O. taurinus and O. oophagus) for the 16S gene fragment used in amphibian DNA barcoding. We propose a new taxonomy, securing the monophyly of Osteocephalus and Tepuihyla by rearranging and redefining the content of both genera and also erect a new genus for the sister group of Osteocephalus. The colouration of newly metamorphosed individuals is proposed as a morphological synapomorphy for Osteocephalus. We recognize and define five monophyletic species groups within Osteocephalus, synonymize three species of Osteocephalus (O. germani, O. phasmatus and O. vilmae) and three species of Tepuihyla (T. celsae, T. galani and T. talbergae) and reallocate three species (Hyla helenae to Osteocephalus, O. exophthalmus to Tepuihyla and O. pearsoni to Dryaderces gen. n.). Furthermore, we flag nine putative new species (an increase to 138% of the current diversity). We conclude that species numbers are largely underestimated, with most hidden diversity centred on widespread and polymorphic nominal species. The evolutionary origin of breeding strategies within Osteocephalus is discussed in the light of this new phylogenetic hypothesis, and a novel type of amplexus (gular amplexus) is described.

SPECIES IDENTITY IN THE GENUS ADENOMERA (ANURA: LEPTODACTYLIDAE) IN SOUTHEASTERN PERU

The genus Adenomera has been a difficult group for systematic studies because the species are similar and geographically variable. Two species have been reported from the Peruvian Amazon Basin: Adenomera andreae and Adenomera hylaedactyla. However, acoustic recordings from the Tambopata National Reserve in southeastern Peru reveal four sympatric advertisement call types that are distinctive in acoustic parameters and to the human ear. Some subtle morphological differences are also present. We conclude that there are at least four sympatric species at Tambopata and that Adenomera has a greater species diversity than currently acknowledged.

Nature conservation: priority-setting needs a global change

The limited resources available for the conservation of biodiversity and ecosystem services call for prioritisation schemes. For instance, in the process of systematic conservation planning site selection is partly determined by efficiency gains. In this paper we present an alternative method for global spatial priority-setting based on ecological indicators, combined with social and economic conditions that influence the effectiveness of conservation, and measures for the long-term persistence of biodiversity. In the analysis the assumption made is that nature conservation should prioritize the effective maintenance of functional ecosystems that do not only provide the most ecosystem services but are also more likely to have a high adaptive capacity towards unavoidable environmental change. Furthermore, the effectiveness and permanence of conservation projects is tied to certain socioeconomic and political conditions that, as we suggest, should be evaluated as part of the conservation priority-setting process. We propose three new priority categories: eco-functionally wise (EcoWise), socioeconomically wise (SocioWise) and proactive allocation of conservation resources considering future climate change (ClimateWise) expressed as indices based on 16 different indicators. Analysing the combined effects of these three categories (EcoSocioClimateWise), in a spatially explicit way highlights the importance of tropical, subtropical but also some temperate and boreal forest areas all of which are characterized by high values of vegetation density, tree height and carbon storage. Our recommendations for policy makers prompt a shift in conservation planning towards advocating the use of ecological and socioeconomic indicators in combination with proxies for the vulnerability to future climate change impacts.

The Influence of Climatic Seasonality on the Diversity of Different Tropical Pollinator Groups

Tropical South America is rich in different groups of pollinators, but the biotic and abiotic factors determining the geographical distribution of their species richness are poorly understood. We analyzed the species richness of three groups of pollinators (bees and wasps, butterflies, hummingbirds) in six tropical forests in the Bolivian lowlands along a gradient of climatic seasonality and precipitation ranging from 410 mm to 6250 mm. At each site, we sampled the three pollinator groups and their food plants twice for 16 days in both the dry and rainy seasons. The richness of the pollinator groups was related to climatic factors by linear regressions. Differences in species numbers between pollinator groups were analyzed by Wilcoxon tests for matched pairs and the proportion in species numbers between pollinator groups by correlation analyses. Species richness of hummingbirds was most closely correlated to the continuous availability of food, that of bees and wasps to the number of food plant species and flowers, and that of butterflies to air temperature. Only the species number of butterflies differed significantly between seasons. We were not able to find shifts in the proportion of species numbers of the different groups of pollinators along the study gradient. Thus, we conclude that the diversity of pollinator guilds is determined by group-specific factors and that the constant proportions in species numbers of the different pollinator groups constitute a general pattern.

A New Species of the discoidalis Group of Eleutherodactylus (Anura, Leptodactylidae) from Inner-Andean Dry Valleys of Bolivia

A new species of the Eleutherodactylus discoidalis group is described from the inner-Andean dry valleys of Bolivia. The new species is similar to Eleutherodactylus cruralis but differs from it in having the discs of fingers III and IV two times wider than the digits proximal to the discs, and the tympanum length being more than half the eye length. Moreover, the advertisement call of the new species is much shorter, has a lower dominant frequency, and fewer notes per call.

The stream-dwelling tadpole of Hyloscirtus charazani (Anura: Hylidae) from Andean Bolivia

Abstract We describe the lotic larva of Hyloscirtus charazani from Charazani and nearby areas, Departamento La Paz, Bolivia (2700–3200 m a.s.l.). It is morphologically undistinguishable from tadpoles of H. armatus sensu lato. These tadpoles show stream adaptation through a robust body with strong tail musculature, enlarged oral disc, increased number of labial tooth rows, and complete marginal papillae. Furthermore, they have a large saccular structure that underlies the limb buds, encloses the vent tube, and partially covers the hindlimbs during their development. Resumen Describimos la larva lótica de Hyloscirtus charazani de Charazani y de otra localidad cercana, Departamento La Paz, Bolivia (2700–3200 m sobre el nivel del mar). Morfologicamente es identica con renacuajos de H. armatus sensu lato. Estos renacuajos muestran caracteres comunmente entendidos como adaptaciones a la vida en arroyos rapidos, como ser el cuerpo robusto, musculatura de la cola robusta, disco oral ensanchado, un aumento en las filas de dientes labiales y papila marginal completa. Ademas, tienen una estructura sacular grande, bajo los primordios de los miembros posteriores, que contiene al tubo cloacal, y que envuelve parcialmente a los miembros durante su desarrollo.

Predicting Leptodactylus (Amphibia, Anura, Leptodactylidae) Distributions: Broad-Ranging Versus Patchily Distributed Species Using a Presence-Only Environmental Niche Modeling Technique

ABSTRACT. Locality data available for many, if not most, species of Neotropical frogs are based on written descriptions of the collecting sites, not on GPS device determined coordinate data. The pre-GPS device data are imprecise relative to GPS data. Niche modeling is a powerful technique for predicting geographic distributions that provides the best results when the locality data are precise. The purpose of this study is to determine whether imprecise historical locality data are sufficient such that niche modeling techniques can yield realistic new insights to species-level distributions. Two sets of frogs of the genus Leptodactylus that have known different kinds of distributions are evaluated: two species with broad, presumably continuous distributions, and four species known to occur in patchy, disjunct habitats in South America. BIOCLIM, a presence-only environmental niche modeling algorithm,was used to define suitable occupancy areas based on multiple sets of environmental parameters that include: monthly mean, max, and min temperatures, and monthly precipitation. A Nature Conservancy — Natureserve ecoregion layer and a high resolution elevation layer were also included in the analyses. Our analyses yield new realistic insights and questions regarding distributions of the Leptodactylus species we evaluated. We recommend incorporation of the Nature Conservancy — Natureserve layer to evaluate Neotropical distributions, as the layer gave much more robust results than use of only the climatic variable analyses.

New Species of Ischnocnema (Anura: Leptodactylidae) from the Andes of Bolivia

Abstract A new species of Ischnocnema is described from cloud forests in the Andes of northwestern Bolivia (La Paz Department). The new species can be distinguished from all other Ischnocnema except Ischnocnema sanctaecrucis by the presence of a pale sacral stripe from the level of the caudal tip of the coccyx to the level of the sacrum, well-developed occipital and sacral folds, rounded narrow terminal discs 1.3 times as wide as the corresponding phalange, and dorsal skin moderately warty. It can be distinguished from I. sanctaecrucis, also of the Bolivian cloud forests, by having head longer than wide, well-marked occipital and sacral folds, longer subacuminate to rounded snout, ventral regions mostly cream with little brown mottling, and dorsal region without reddish to scarlet mottling or spots. The new species shares some characteristics with other species of Ischnocnema and members of the Eleutherodactylus discoidalis group, and it appears closely related to I. sanctaecrucis.

Another New Species of Small Hyla (Anura, Hylidae) from Amazonian Sub-Andean Forest of Western Bolivia

Abstract We describe a new species of small Hyla from sub-Andean forest of the Departamento La Paz, Bolivia. The new species is mainly characterized by smooth dorsal skin with few minute scattered tubercles, presence of a tympanic annulus, lack of tarsal folds, a brown dorsum with irregular longitudinal dark brown stripes, a broad dark brown lateral stripe extending to loreal region, and advertisement call. The phylogenetic relationships of the new species remain unknown, but given the morphological similarities of adults, we tentatively place it in the Hyla microcephala group.

Priority areas for amphibian conservation in a neotropical megadiverse country: the need for alternative, non place based, conservation

Amphibians constitute one of the most threatened species group worldwide. Because they are affected by various threats such as habitat destruction, chytrid fungus, climate change, invasive species and human use, different approaches are needed for their conservation. In this paper we examine patterns of alpha diversity, endemism and threatened species distributions of amphibians in Bolivia. We modeled distribution data using ecological niche modeling implemented with MaxEnt (Maximum Entropy Modeling). In contrast with previous studies, we validated each individual map with specialists. Important areas for amphibian conservation were defined by overlapping species richness, endemism richness and concentration of threatened species, and then excluding all unsuitable habitats. Our goal was to identify high-priority sites for the conservation of amphibians in Bolivia. We discuss a place-based strategy (identification of priority areas) and the need for further non place-based strategies. We highlight the importance of strengthening two existing protected areas that together host up to 40% of Bolivia’s threatened amphibian species, and we propose alternative strategies such as ex situ conservation for those species not covered by the priority areas. Bolivia’s General Directorate for Biodiversity and Protected Areas (in Spanish, Dirección General de Biodiversidad y Áreas Protegidas (DGBAP)) and Bolivian experts are jointly using the here presented results to define a national strategy of amphibian conservation for safeguarding this important element of Bolivia’s biodiversity.