No abstract

Biogeographical regions (bioregions) reveal how different sets of species are spatially grouped and therefore are important units for conservation, historical biogeography, ecology, and evolution. Several methods have been developed to identify bioregions based on species distribution data rather than expert opinion. One approach successfully applies network theory to simplify and highlight the underlying structure in species distributions. However, this method lacks tools for simple and efficient analysis. Here, we present Infomap Bioregions, an interactive web application that inputs species distribution data and generates bioregion maps. Species distributions may be provided as georeferenced point occurrences or range maps, and can be of local, regional, or global scale. The application uses a novel adaptive resolution method to make best use of often incomplete species distribution data. The results can be downloaded as vector graphics, shapefiles, or in table format. We validate the tool by processing large data sets of publicly available species distribution data of the world’s amphibians using species ranges, and mammals using point occurrences. We then calculate the fit between the inferred bioregions and WWF ecoregions. As examples of applications, researchers can reconstruct ancestral ranges in historical biogeography or identify indicator species for targeted conservation.

The present study is a synthesis on snake diversity and distribution in the Caatinga region of northeastern Brazil, providing an updated species list and data on natural history and geographic distribution. Our study is based on the careful revision of 7,102 voucher specimens, housed in 17 herpetological collections, complemented by data on taxonomic literature. We recorded a total of 112 snake species in the Caatinga, belonging to nine families: Anomalepididae, Leptotyphlopidae, Typhlopidae, Aniliidae, Boidae, Viperidae, Elapidae, Colubridae, and Dipsadidae. Our list includes at least 13 never recorded species for this region, as well as distribution records for all species known from the Caatinga (including expansion and new records of distribution). The snake assemblage of the Caatinga is complex, sharing species with other continental open areas (38.4%), forested areas (27.7%), and both open and forested areas (32.1%). The richest areas were isolated plateaus, followed by contact areas, semi-arid caatinga, and sandy dunes of the São Franscisco River. We identified 22 Caatinga endemic species with the sandy dunes of São Franscico River showing the highest endemism level (12 species, with six endemic species restricted to the area) followed by semi-arid caatinga, and isolated plateaus (eight endemic species each, and six and three endemic species with restricted distribution to each area, respectively). Most species show relatively restricted ranges in parts of the Caatinga. The snake assemblage in Caatinga includes mainly terrestrial species (38.4%), followed by fossorial/cryptozoic (26.8%), arboreal/semi-arboreal (26.8%), and aquatic/semi-aquatic (7.1%) species. Vertebrates are the most important dietary item (80.4%), with 56.6% of species being generalist consumers of this kind of prey; 24.4% are frog-eaters, 7.8% prey on caecilians/amphisbaenians, 6.7% lizard-eaters, 3.3% mammal-eaters, and 1.1% are fish-eaters. Only 18.7% of the snakes eat invertebrate prey, as arthropods, annelids, and mollusks. In relation to time of activity, 35.7% of snakes are both diurnal and nocturnal, 33.0% are strictly nocturnal, and 30.4% are diurnal. The data provided herein increase the list of Caatinga snake species from 50 to 112, and includes detailed maps and information on geographic distribution. The Caatinga snake assemblage shows high richness and endemism levels, and our results highlight the usefulness of basic natural history data and revision of voucher specimens as baseline information for biogeographic studies and conservation strategies.

The unparalleled biodiversity found in the American tropics (the Neotropics) has attracted the attention of naturalists for centuries. Despite major advances in recent years in our understanding of the origin and diversification of many Neotropical taxa and biotic regions, many questions remain to be answered. Additional biological and geological data are still needed, as well as methodological advances that are capable of bridging these research fields. In this review, aimed primarily at advanced students and early-career scientists, we introduce the concept of “trans-disciplinary biogeography,” which refers to the integration of data from multiple areas of research in biology (e.g., community ecology, phylogeography, systematics, historical biogeography) and Earth and the physical sciences (e.g., geology, climatology, palaeontology), as a means to reconstruct the giant puzzle of Neotropical biodiversity and evolution in space and time. We caution against extrapolating results derived from the study of one or a few taxa to convey general scenarios of Neotropical evolution and landscape formation. We urge more coordination and integration of data and ideas among disciplines, transcending their traditional boundaries, as a basis for advancing tomorrow’s ground-breaking research. Our review highlights the great opportunities for studying the Neotropical biota to understand the evolution of life.

MotivationWe generated a novel database of Neotropical snakes (one of the world's richest herpetofauna) combining the most comprehensive, manually compiled distribution dataset with publicly available data. We assess, for the first time, the diversity patterns for all Neotropical snakes as well as sampling density and sampling biases.Main types of variables containedWe compiled three databases of species occurrences: a dataset downloaded from the Global Biodiversity Information Facility (GBIF), a verified dataset built through taxonomic work and specialized literature, and a combined dataset comprising a cleaned version of the GBIF dataset merged with the verified dataset.Spatial location and grainNeotropics, Behrmann projection equivalent to 1° × 1°.Time periodSpecimens housed in museums during the last 150 years.Major taxa studiedSquamata: Serpentes.Software formatGeographical information system (GIS).ResultsThe combined dataset provides the most comprehensive distribution database for Neotropical snakes to date. It contains 147,515 records for 886 species across 12 families, representing 74% of all species of snakes, spanning 27 countries in the Americas. Species richness and phylogenetic diversity show overall similar patterns. Amazonia is the least sampled Neotropical region, whereas most well‐sampled sites are located near large universities and scientific collections. We provide a list and updated maps of geographical distribution of all snake species surveyed.Main conclusionsThe biodiversity metrics of Neotropical snakes reflect patterns previously documented for other vertebrates, suggesting that similar factors may determine the diversity of both ectothermic and endothermic animals. We suggest conservation strategies for high‐diversity areas and sampling efforts be directed towards Amazonia and poorly known species.

Aim Our aims were to test the predictions of the vicariance model, searching for natural, non-random biogeographical units using data on snake distributions, and to assess the conservation of biogeographical patterns and underlying processes in the poorly studied Caatinga region.Location Caatinga region, north-eastern Brazil.Methods We revised and georeferenced 7352 snake occurrence records at point localities, by direct examination of voucher specimens in zoological collections and revision of literature data. We tested two predictions of the vicariance model via biotic element analysis using two datasets (all taxa and endemics) mapped onto a 1°9 1°square grid across the Caatinga. Finally, we examined the overlap between recovered biogeographical units and spatial patterns of habitat loss and protected area coverage. ResultsWe recorded 112 snake species from the Caatinga, of which 22 (20%) are endemics. The predictions of the vicariance model were corroborated by the detection of groups of species with significantly clustered ranges (biotic elements). The analysis with the full dataset detected eight biotic elements, and three endemic biotic elements were found when only using endemics. The three endemic biotic elements correspond to core areas of biotic elements detected with the larger dataset. The average habitat loss for species forming biotic elements was 46%, and was similar among biotic elements. Protected area coverage is different for species from different biotic elements, and most species' ranges are very poorly represented in protected areas.Main conclusions The Caatinga harbours a peculiar snake fauna with significantly clustered species ranges concordant with the predictions of the vicariance model. Our results, representing the first formal test of vicariance patterns in the Caatinga, detected poor overlap between biotic elements and protected areas, indicating that biogeographical patterns and processes are largely unprotected in this imperilled and neglected Neotropical region.

Based on the literature, we had predicted that the diversification within the Neotropical snake genus Bothrops occurred along a latitudinal gradient from north to south, with diversification into unoccupied niches through ecological opportunity, not correlated with geoclimatic events. Using a dated phylogeny and estimating likelihoods of ancestral states at cladogenesis events, we reconstructed ancestral areas and assessed major events of the diversification of Bothrops clades, and we also discuss systematic implications for this group. Based on the phylogeny we produced, B. lojanus was not considered as part of the genus Bothrops since the results recovered this species nested within the Bothrocophias clade. We infer that the diversification of the Miocene Bothrops pictus and Bothrops alternatus clades may be related to the uplift of the western slopes of the Andes and the Argentinian Patagonian Andes, respectively. The Pliocene Bothrops taeniatus and Bothrops osbornei clades may be related to the uplift of the eastern and northern Andes, respectively. The Plio‐Pleistocene Bothrops neuwiedi clade may be related to the habitat transitions from a warmer and forested environment to a cooler and open landscape; the Bothrops jararaca (i.e. island endemic species) and Bothrops lanceolatus clades to over‐water dispersal with island speciation; and Bothrops atrox clade to the appearance of the Panamanian land bridge. We found that a multitemporal and multidirectional history of diversification may be correlated with geoclimatic and dispersalist events. We argue that the vacant niche hypothesis by itself does not explain Bothrops diversification.

Factors driving the spatial configuration of centres of endemism have long been a topic of broad interest and debate. Due to different eco-evolutionary processes, these highly biodiverse areas may harbour different amounts of ancient and recently diverged organisms (paleo-and neo-endemism, respectively). Patterns of endemism still need to be measured at distinct phylogenetic levels for most clades and, consequently, little is known about the distribution, the age and the causes of such patterns. Here we tested for the presence of centres with high phylogenetic endemism (PE) in the highly diverse Neotropical snakes, testing the age of these patterns (paleo-or neoendemism), and the presence of PE centres with distinct phylogenetic composition. We then tested whether PE is predicted by topography, by climate (seasonality, stability, buffering and relictualness), or biome size. We found that most areas of high PE for Neotropical snakes present a combination of both ancient and recently diverged diversity, which is distributed mostly in the Caribbean region, Central America, the Andes, the Atlantic Forest and on scattered highlands in central Brazil. Turnover of lineages is higher across Central America, resulting in more phylogenetically distinct PE centres compared to South America, which presents a more phylogenetically uniform snake Research * equally contributed. 329fauna. Finally, we found that elevational range (topographic roughness) is the main predictor of PE, especially for paleoendemism, whereas low paleo-endemism levels coincide with areas of high climatic seasonality. Our study highlights the importance of mountain systems to both ancient and recent narrowly distributed diversity. Mountains are both museums and cradles of snake diversity in the Neotropics, which has important implications for conservation in this region.