Co-occurrence patterns between false coral snake Atractus latifrons (Günther, 1868) (Serpentes: Dipsadidae) and venomous coral snakes from the Amazon.

Batesian mimicry may result in remarkable cases of phenotypic convergence that represent classic examples of evolution through natural selection. The existence of mimicry systems among coral snakes, however, remains controversial because of contradictions between the predictions of mimetic theory and the empirical patterns of co-occurrence and species abundance. Here, we analyze the geographic distribution of coral snake species of the genus Micrurus and populations of the false coral snake Atractus latifrons in Amazonia, and perform ecological niche modeling (ENM) analyzes to generate potential geographic distributions of species of Micrurus and A. latifrons, identify patterns of co-occurrence and assess whether the distribution of A. latifrons coincides with the distribution of Micrurus species, which could suggest the existence of a possible mimetic relationship between the species. We identified six Micrurus species that may represent mimetic models for A. latifrons. The results of the co-occurrence analysis corroborates the results from ENM, indicating that chromatic patterns of A. latifrons and their respective model species are aggregated. Our study suggests that all color patterns of A. latifrons - including the tricolor monads, and the more common tricolor dyads and tricolor tetrads - may benefit from the resemblance with other Micrurus species as perfect and imperfect mimics.

Back home? Uncertainties for returning seized animals to the source-areas under climate change.

Regardless of the economic, social and environmental impacts caused by wild animal trafficking worldwide, the suitable destination of seized specimens is one of the main challenges faced by environmental managers and authorities. In Brazil, returning seized animals to the wild has been the most frequent path in population restoration programs, and has been carried out, as a priority, in areas where the animals were captured. However, in addition to the difficulty in identifying the locations of illegal captures, little scientific knowledge is available on the future viability of the source-areas to global climate change. Thus, the current work aims to evaluate the impacts of climate change on the main source-municipalities for animal trafficking in Brazil, referred to herein as source-areas. For this, using ecological niche modeling, the environmental suitability of the source-areas for illegal animal captures was evaluated in two scenarios at two different time horizons: optimistic (RCP 26) and pessimistic (RCP 85) emission scenarios in both 2050 and 2070 projections. Moreover, the source-areas were compared with the Brazilian Federal protected areas, used here as the control group. According to the results, Brazilian source-municipalities are not always the best option for maintaining the most seized species in the future simulations, and, therefore, seem not be the best option for projects that aim for the return of these animals to the wild. In this sense, despite the genetic and ecological issues inherent in translocation projects, our results suggest that population restoration programs for seized species need to be rethought, and furthermore other suitable areas could be considered for truly ensuring the survival and maintenance of overexploited populations in the long term.

Demographical history and palaeodistribution modelling show range shift towards Amazon Basin for a Neotropical tree species in the LGM.

BACKGROUND: We studied the phylogeography and demographical history of Tabebuia serratifolia (Bignoniaceae) to understand the disjunct geographical distribution of South American seasonally dry tropical forests (SDTFs). We specifically tested if the multiple and isolated patches of SDTFs are current climatic relicts of a widespread and continuously distributed dry forest during the last glacial maximum (LGM), the so called South American dry forest refugia hypothesis, using ecological niche modelling (ENM) and statistical phylogeography. We sampled 235 individuals of T. serratifolia in 17 populations in Brazil and analysed the polymorphisms at three intergenic chloroplast regions and ITS nuclear ribosomal DNA. RESULTS: Coalescent analyses showed a demographical expansion at the last c. 130 ka (thousand years before present). Simulations and ENM also showed that the current spatial pattern of genetic diversity is most likely due to a scenario of range expansion and range shift towards the Amazon Basin during the colder and arid climatic conditions associated with the LGM, matching the expected for the South American dry forest refugia hypothesis, although contrasting to the Pleistocene Arc hypothesis. Populations in more stable areas or with higher suitability through time showed higher genetic diversity. Postglacial range shift towards the Southeast and Atlantic coast may have led to spatial genome assortment due to leading edge colonization as the species tracks suitable environments, leading to lower genetic diversity in populations at higher distance from the distribution centroid at 21 ka. CONCLUSION: Haplotype sharing or common ancestry among populations from Caatinga in Northeast Brazil, Atlantic Forest in Southeast and Cerrado biome and ENM evince the past connection among these biomes.

Conservation biogeography of the Cerrado's wild edible plants under climate change: Linking biotic stability with agricultural expansion.

UNLABELLED: REMISE OF THE STUDY: Wild edible plants (WEPs) have an important cultural and economic role in human population worldwide. Human impacts are quickly converting natural habitats in agricultural, cattle ranch, and urbanized lands, putting native species on peril of risk of extinction, including some WEPs. Moreover, global climate changes also can pose another threat to species persistency. Here, we established conservation priorities for the Cerrado, a neotropical region in South America with high levels of plant endemism and vulnerability, aiming to assure long-term persistency of 16 most important WEPs. We evaluated these conservation priorities using a conservation biogeography framework using ecological patterns and process at a biogeographical scale to deal with species conservation features. METHODS: We built ecological niche models for 16 WEPs from Cerrado in the neotropics using climate models for preindustrial, past (Last Glacial Maximum) and future (year 2080) time periods to establish climatically stable areas through time, finding refugias for these WEPs. We used a spatial prioritization algorithm based on the spatial pattern of irreplaceability across the neotropics, aiming to ensure the persistence of at least 25% of range size in climatically stable areas for each WEP, using agricultural models as constraints. KEY RESULTS: The Southeast Cerrado was the most biotically stable and irreplaceable region for the WEPs compared with other areas across the neotropics. CONCLUSIONS: Our findings strongly suggest that the Southeast Cerrado should be considered a conservation priority, with new protected areas to be sustainably managed and restored, to guarantee the supply of cultural and ecosystem services provided from the Cerrado's WEPs.

Recovering species demographic history from multi-model inference: the case of a Neotropical savanna tree species.

BACKGROUND: Glaciations were recurrent throughout the Quaternary and potentially shaped species genetic structure worldwide by affecting population dynamics. Here, we implemented a multi-model inference approach to recover the distribution dynamics and demographic history of a Neotropical savanna tree, Tabebuia aurea (Bignoniaceae). Exploring different algorithms and paleoclimatic simulations, we used ecological niche modelling to generate alternative hypotheses of potential demographic changes through the last glacial cycle and estimated genetic parameters using coalescent modelling. RESULTS: Comparing predictions from demographic hypotheses with genetic parameters of modern populations, our findings revealed a likely scenario of population decline, with spatial displacement towards Northeast Brazil from the last glacial maximum to the mid-Holocene. Subsequently, populations expanded in response to the return of the climatically suitable conditions in Central-West Brazil. Nevertheless, a wide historical refugium across Central Brazil likely maintained large populations connected throughout time. The expected genetic signatures from such predicted distribution dynamics are also corroborated by spatial genetic structure observed in modern populations. CONCLUSION: By exploring uncertainties inherent in multiple working hypotheses, we have shown that multi-model inference is a fruitful and efficient approach to recover the nature, timing and geographical context of the Tabebuia aurea population dynamic in response to the Quaternary climate changes.

No evidence for niche competition in the extinction of the South American saber-tooth species.

The end of South American isolation during the Great American Biotic Interchange (GABI) promoted the contact between South and North American saber-tooth forms that evolved in isolation. This contact may have driven saber-tooth species to a competitive interaction, resulting in the extinction of the South American saber-tooth form. Here, we used paleoclimatic data to compare the climatic niche of the saber-tooth forms Thylacosmilus atrox (from South America), Smilodon fatalis, and Smilodon populator (both originally from North America). We evaluated niche width, overlap, and similarity to infer potential geographic distribution overlap and competition between these North and South American predators. To do so, we obtained the climatic variables from sites where occurrence fossil records were available. Our results suggest that T. atrox had a narrower climatic niche compared to Smilodon species. Although we found a significant climatic niche overlap and similarity between S. fatalis and T. atrox, it seems unlikely that both species have co-occurred. Low niche overlap and similarity between T. atrox and S. populator dismiss competitive interaction between them. Moreover, climatic niche and low tolerance for environmental changes may have been the cause of the South American saber-tooth extinction.

A coupled phylogeographical and species distribution modelling approach recovers the demographical history of a Neotropical seasonally dry forest tree species.

We investigated here the demographical history of Tabebuia impetiginosa (Bignoniaceae) to understand the dynamics of the disjunct geographical distribution of South American seasonally dry forests (SDFs), based on coupling an ensemble approach encompassing hindcasting species distribution modelling and statistical phylogeographical analysis. We sampled 17 populations (280 individuals) in central Brazil and analysed the polymorphisms at chloroplast (trnS-trnG, psbA-trnH, and ycf6-trnC intergenic spacers) and nuclear (ITS nrDNA) genomes. Phylogenetic analyses based on median-joining network showed no haplotype sharing among population but strong evidence of incomplete lineage sorting. Coalescent analyses showed historical constant populations size, negligible gene flow among populations, and an ancient time to most recent common ancestor dated from ~4.7 ± 1.1 Myr BP. Most divergences dated from the Lower Pleistocene, and no signal of important population size reduction was found in coalescent tree and tests of demographical expansion. Demographical scenarios were built based on past geographical range dynamic models, using two a priori biogeographical hypotheses ('Pleistocene Arc' and 'Amazonian SDF expansion') and on two additional hypotheses suggested by the palaeodistribution modelling built with several algorithms for distribution modelling and palaeoclimatic data. The simulation of these demographical scenarios showed that the pattern of diversity found so far for T. impetiginosa is in consonance with a palaeodistribution expansion during the last glacial maximum (LGM, 21 kyr BP), strongly suggesting that the current disjunct distribution of T. impetiginosa in SDFs may represent a climatic relict of a once more wide distribution.

Global richness patterns of venomous snakes reveal contrasting influences of ecology and history in two different clades.

Recent studies addressing broad-scale species richness gradients have proposed two main primary drivers: contemporary climate and evolutionary processes (differential balance between speciation and extinction). Here, we analyze the global richness patterns of two venomous snake clades, Viperidae and Elapidae. We used ordinary least squares multiple regression (OLS) and partial regression analysis to investigate to what extent actual evapotranspiration (AET; summarizing current environmental conditions) and biogeographical regions (representing evolutionary effects) were associated with species richness. For viperids, AET explained 45.6% of the variance in richness whereas the effect of this variable for elapids was almost null (0.5%). On the other hand, biogeographic regions were the best predictors of elapid richness (56.5%), against its relatively small effect (25.9%) in viperid richness. Partial regressions also revealed similar patterns for independent effects of climate and history in both clades. However, the independent historical effect in Elapidae decreased from 45.2 to 17.8% when we excluded Australia from the analyses, indicating that the strong historical effect that had emerged for the global richness pattern was reflecting the historical process of elapid radiation into Australia. Even after excluding Australia, the historical signal in elapid richness in the rest of the globe was still significant and much higher than that observed in viperid richness at a global scale (2.7% after controlling for AET effects). Differences in the evolutionary age of these two clades can be invoked to explain these contrasting results, in that viperids probably had more time for diversification, generating richness responses to environmental gradients, whereas the pattern of distribution of elapid richness can be more directly interpreted in an evolutionary context. Moreover, these results show the importance of starting to adopt deconstructive approaches to species richness, since the driving factors of these patterns may vary from group to group according to their evolutionary history.

Historical range contractions can predict extinction risk in extant mammals.

Climate change is amongst the main threats to biodiversity. Considering extant mammals endured Quaternary climate change, we analyzed the extent to which this past change predicts current mammals' extinction risk at global and biogeographical scales. We accessed range dynamics by modeling the potential distribution of all extant terrestrial mammals in the Last Glacial Maximum (LGM, 21,000 years ago) and in current climate conditions and used extinction risk from IUCN red list. We built General Linear Mixed-Effects Models to test the magnitude with which the variation in geographic range (ΔRange) and a proxy for abundance (ΔSuitability) between the LGM and present-day predicts current mammal's extinction risk. We found past climate change most strongly reduced the geographical range and climatic suitability of threatened rather than non-threatened mammals. Quaternary range contractions and reduced suitability explain around 40% of species extinction risk, particularly for small-bodied mammals. At global and biogeographical scales, all groups that suffered significant Quaternary range contractions now contain a greater proportion of threatened species when compared to groups whose ranges did not significantly contract. This reinforces the importance of using historical range contractions as a key predictor of extinction risk for species in the present and future climate change scenarios and supports current efforts to fight climate change for biodiversity conservation.

Climate change will decrease the range size of snake species under negligible protection in the Brazilian Atlantic Forest hotspot.

Reptiles are highly susceptible to climate change, responding negatively to thermal and rainfall alterations mainly in relation to their reproductive processes. Based on that, we evaluated the effects of climate change on climatically suitable areas for the occurrence of snakes in the Atlantic Forest hotspot, considering the responses of distinct reproductive groups (oviparous and viviparous). We assessed the species richness and turnover patterns affected by climate change and projected the threat status of each snake species at the end of the century. We also evaluated the effectiveness of the protected areas in safeguarding the species by estimating the mean percentage overlap between snake species distribution and protected areas (PAs) network and by assessing whether such areas will gain or lose species under climate change. Our results showed greater species richness in the eastern-central portion of the Atlantic Forest at present. In general, we evidenced a drastic range contraction of the snake species under climate change. Temporal turnover tends to be high in the western and north-eastern edges of the biome, particularly for oviparous species. Our predictions indicate that 73.6% of oviparous species and 67.6% of viviparous species could lose at least half of their original range by 2080. We also found that existing protected areas of the Atlantic Forest Hotspot have a very limited capacity to safeguard snakes at the current time, maintaining the precarious protection in the future, with the majority of them predicted to lose species at the end of this century. Although oviparous and viviparous snakes have been designated to be dramatically impacted, our study suggests a greater fragility of the former in the face of climate change. We advocated that the creation of new protected areas and/or the redesign of the existing network to harbour regions that maximize the snake species occupancy in the face of future warming scenarios are crucial measures for the conservation of this group.

Reducing Wallacean shortfalls for the coralsnakes of the Micrurus lemniscatus species complex: Present and future distributions under a changing climate.

South American coralsnakes are characterized by inconspicuous and poorly known species, which are potentially very sensitive to climate change. Here, we assess the impact of future climate change on the distributions of the Micrurus lemniscatus species complex after addressing the Wallacean shortfalls and refining the knowledge about their current geographic distributions. We also evaluate the efficiency of the current reserve network to protect the species in the present and future. We applied ecological niche model tools through a carefully examined set of occurrence records to generate potential present distributions and to project these distributions into future scenarios of climate change. Specific thresholds based on occurrence records along with expert opinions were used to delineate the geographic distribution of each species. A hierarchical ANOVA was applied to evaluate the uncertainties in species distributions across niche modeling methods and climate models and nested into the time factor (present and future). Multiple regression models were used to infer the relative importance of the climatic variables to determine the species' suitability. A gap analysis was performed to address the representativeness of species distributions into protected areas. Predicted geographic distributions were compatible with the known distributions and the expert opinions, except for M. l. carvalhoi. New areas for field research were identified. Variation in precipitation was the most important factor defining the habitat suitability for all species, except for M. diutius. All taxa (except M. l. lemniscatus) will shrink their distributions in the future; less than 50% of the present suitable areas are protected in reserve networks, and less than 40% of these areas will be held in reserves in the future. We found strong evidence that coralsnakes may be highly sensitive to the ongoing changes and must be protected.

Coalescent Simulation and Paleodistribution Modeling for Tabebuia rosealba Do Not Support South American Dry Forest Refugia Hypothesis.

Studies based on contemporary plant occurrences and pollen fossil records have proposed that the current disjunct distribution of seasonally dry tropical forests (SDTFs) across South America is the result of fragmentation of a formerly widespread and continuously distributed dry forest during the arid climatic conditions associated with the Last Glacial Maximum (LGM), which is known as the modern-day dry forest refugia hypothesis. We studied the demographic history of Tabebuia rosealba (Bignoniaceae) to understand the disjunct geographic distribution of South American SDTFs based on statistical phylogeography and ecological niche modeling (ENM). We specifically tested the dry forest refugia hypothesis; i.e., if the multiple and isolated patches of SDTFs are current climatic relicts of a widespread and continuously distributed dry forest during the LGM. We sampled 235 individuals across 18 populations in Central Brazil and analyzed the polymorphisms at chloroplast (trnS-trnG, psbA-trnH and ycf6-trnC intergenic spacers) and nuclear (ITS nrDNA) genomes. We performed coalescence simulations of alternative hypotheses under demographic expectations from two a priori biogeographic hypotheses (1. the Pleistocene Arc hypothesis and, 2. a range shift to Amazon Basin) and other two demographic expectances predicted by ENMs (3. expansion throughout the Neotropical South America, including Amazon Basin, and 4. retraction during the LGM). Phylogenetic analyses based on median-joining network showed haplotype sharing among populations with evidence of incomplete lineage sorting. Coalescent analyses showed smaller effective population sizes for T. roseoalba during the LGM compared to the present-day. Simulations and ENM also showed that its current spatial pattern of genetic diversity is most likely due to a scenario of range retraction during the LGM instead of the fragmentation from a once extensive and largely contiguous SDTF across South America, not supporting the South American dry forest refugia hypothesis.

A Short Guide to the Climatic Variables of the Last Glacial Maximum for Biogeographers.

Ecological niche models are widely used for mapping the distribution of species during the last glacial maximum (LGM). Although the selection of the variables and General Circulation Models (GCMs) used for constructing those maps determine the model predictions, we still lack a discussion about which variables and which GCM should be included in the analysis and why. Here, we analyzed the climatic predictions for the LGM of 9 different GCMs in order to help biogeographers to select their GCMs and climatic layers for mapping the species ranges in the LGM. We 1) map the discrepancies between the climatic predictions of the nine GCMs available for the LGM, 2) analyze the similarities and differences between the GCMs and group them to help researchers choose the appropriate GCMs for calibrating and projecting their ecological niche models (ENM) during the LGM, and 3) quantify the agreement of the predictions for each bioclimatic variable to help researchers avoid the environmental variables with a poor consensus between models. Our results indicate that, in absolute values, GCMs have a strong disagreement in their temperature predictions for temperate areas, while the uncertainties for the precipitation variables are in the tropics. In spite of the discrepancies between model predictions, temperature variables (BIO1-BIO11) are highly correlated between models. Precipitation variables (BIO12-BIO19) show no correlation between models, and specifically, BIO14 (precipitation of the driest month) and BIO15 (Precipitation Seasonality (Coefficient of Variation)) show the highest level of discrepancy between GCMs. Following our results, we strongly recommend the use of different GCMs for constructing or projecting ENMs, particularly when predicting the distribution of species that inhabit the tropics and the temperate areas of the Northern and Southern Hemispheres, because climatic predictions for those areas vary greatly among GCMs. We also recommend the exclusion of BIO14 and BIO15 from ENMs because those variables show a high level of discrepancy between GCMs. Thus, by excluding them, we decrease the level of uncertainty of our predictions. All the climatic layers produced for this paper are freely available in http://ecoclimate.org/.

Correlation between genetic diversity and environmental suitability: taking uncertainty from ecological niche models into account.

The hindcast of shifts in the geographical ranges of species as estimated by ecological niche modelling (ENM) has been coupled with phylogeographical patterns, allowing the inference of past processes that drove population differentiation and genetic variability. However, more recently, some studies have suggested that maps of environmental suitability estimated by ENM may be correlated to species' abundance, raising the possibility of using environmental suitability to infer processes related to population demographic dynamics and genetic variability. In both cases, one of the main problems is that there is a wide variation in ENM development methods and climatic models. In this study, we analyse the relationship between heterozygosity (He) and environmental suitability from multiple ENMs for 25 population estimates for Dipteryx alata, a widely distributed, endemic tree species of the Cerrado region of central Brazil. We propose a new approach for generating a statistical distribution of correlations under randomly generated ENM. The confidence intervals from these distributions indicate how model selection with different properties affects the ability to detect a correlation of interest (e.g. the correlation between He and suitability). Additionally, our approach allows us to explore which particular ensemble of ENMs produces the better result for finding an association between environmental suitability and He. Caution is necessary when choosing a method or a climatic data set for modelling geographical distributions, but the new approach proposed here provides a conservative way to evaluate the ability of ensembles to detect patterns of interest.

Body size, extinction risk and knowledge bias in New World snakes.

Extinction risk and body size have been found to be related in various vertebrate groups, with larger species being more at risk than smaller ones. We checked whether this was also the case for snakes by investigating extinction risk-body size relationships in the New World's Colubroidea species. We used the IUCN Red List risk categories to assign each species to one of two broad levels of threat (Threatened and Non-Threatened) or to identify it as either Data Deficient or Not-Evaluated by the IUCN. We also included the year of description of each species in our analysis as this could affect the level of threat assigned to it (earlier described species had more time to gather information about them, which might have facilitated their evaluation). Also, species detectability could be a function of body size, with larger species tending to be described earlier, which could have an impact in extinction risk-body size relationships. We found a negative relationship between body size and description year, with large-bodied species being described earlier. Description year also varied among risk categories, with Non-Threatened species being described earlier than Threatened species and both species groups earlier than Data Deficient species. On average, Data Deficient species also presented smaller body sizes, while no size differences were detected between Threatened and Non-Threatened species. So it seems that smaller body sizes are related with species detectability, thus potentially affecting both when a species is described (smaller species tend to be described more recently) as well as the amount of information gathered about it (Data Deficient species tend to be smaller). Our data also indicated that if Data Deficient species were to be categorized as Threatened in the future, snake body size and extinction risk would be negatively related, contrasting with the opposite pattern commonly observed in other vertebrate groups.

Padrões espaciais da riqueza de espécies de viperídeos na Américado Sul: temperatura ambiental vs. cinética-bioquímica / Spatial patterns of viperid species richness in South America: environmental temperature vs. biochemical kinetics

Este estudo tem por objetivo testar a influência da temperatura na taxametabólica de viperídeos como um mecanismo básico de origem dos padrões espaciais deriqueza desse grupo na América do Sul, como proposto por Allen et al. (2002) dentro daTeoria Metabólica em Ecologia. Para isso, testamos a relação entre o logaritmo natural dariqueza de espécies de viperídeos e o inverso da temperatura (em Kelvin, 1000*K-1), apóscorrigir os efeitos da autocorrelação espacial, e verificamos se a reta estimada apresentainclinação de -9,0*T. As variáveis apresentaram baixo índice de correlação (r2 = 0,216; p <0,0001), com uma inclinação da reta de -3,737*T (C.I. (95%) ± 0,379). Os resultadosindicaram que os viperídeos não respondem à variação de temperatura da mesma forma queos demais grupos testados, uma vez que o intervalo de confiança para o ângulo da retaestimada não contempla o valor -9,0*T, como predito pelo modelo. O presente estudosugere que o padrão espacial da riqueza de espécies de viperídeos na América do Sul éestruturado por outros parâmetros além da temperatura, não contemplados no modelo deAllen et al. (2002).

The aim of this study was to testthe influence of temperature on metabolic rates of viperid species as the underlyingmechanism to explain the richness pattern of this group in South America, following theMetabolic Theory of Ecology (MTE) proposed by Allen et al. (2002). We tested MTEpredictions by considering the relationship between the natural logarithm of viperid speciesrichness and the inverse of temperature (in Kelvin, 1000*K-1) after to correct for spatialautocorrelation effects and to check whether the linear function presents a slope of -9.0*T.The relationship between variables presented low correlation coefficient (r2 = 0.216; P <0.0001) and a slope of -3.737*T (C.I. (95%) ± 0.379).These results showed that viperidsrespond in a different way to the temperature gradient in comparison with other taxa andthe prediction of Allen et al. (2002), since the confidence interval of slope in this case doesnot include the value of -9.0*T. This study demonstrates that temperature is not the soledriver of broad-scale spatial pattern of viperid species richness in South America.