Genetic data improve the assessment of the conservation status based only on herbarium records of a Neotropical tree

Although there is a consensus among conservation biologists about the importance of genetic information, the assessment of extinction risk and conservation decision-making generally do not explicitly consider this type of data. Genetic data can be even more important in species where little other information is available. In this study, we investigated a poorly known legume tree, Dimorphandra exaltata, from the Brazilian Atlantic Forest, a hotspot for conservation. We coupled species distribution models and geospatial assessment based on herbarium records with population genetic analyses to evaluate its genetic status and extinction risk, and to suggest conservation measures. Dimorphandra exaltata shows low genetic diversity, inbreeding, and genetic evidence of decrease in population size, indicating that the species is genetically depleted. Geospatial assessment classified the species as Endangered. Species distribution models projected a decrease in range size in the near future (2050). The genetic status of the species suggests low adaptive potential, which compromises its chances of survival in the face of ongoing climatic change. Altogether, our coupled analyses show that the species is even more threatened than indicated by geospatial analyses alone. Thus, conservation measures that take into account genetic data and the impacts of climate change in the species should be implemented.


Results
Genetic status: genetic diversity and structure, migration, and effective population size. Of the 11 microsatellites loci used, only Dw103 for CON population and Dw28 for all populations showed null alleles, while scoring errors due to stuttering or large allele dropout were not found. However, as the genetic diversity estimates were similar between the original dataset and the corrected for null alleles dataset using the Brookfield 1 method in MicroChecker version 2.2.0.2 28 , we considered the estimates without null allele correction. No significant linkage disequilibrium between loci was observed in any comparison after Bonferroni correction. To estimate the genetic diversity and structure of D. exaltata we also included the seven isolated individuals of the Atlantic Forest in addition to the three sampled populations. These individuals from three sites with distances ranging from 31.5 to 95.7 km were analysed together (AFI group; Table 1) to represent the diversity of eastern Atlantic Forest, although they probably do not represent samples of only one population. For the remaining genetic analyses, the AFI group was not included.
Among the three populations and the AFI group, the mean A and A R ranged from 3.091 to 4.545 and from 2.348 to 3.419, respectively. H O and H E per population ranged from 0.320 to 0.481 and from 0.289 to 0.560, respectively, with mean values of H O and H E of 0.372 and 0.413, respectively (Table 1). Populations CON and SER showed the higher values of private alleles with 20 and 16 alleles, respectively. In addition, nine private alleles were found in the isolated individuals from Atlantic Forest, despite the small sample size ( Table 1). The exact test showed that two of the three populations, SER and CON, showed significant positives inbreeding coefficients (F IS ) after Bonferroni correction, with estimated values of 0.195 and 0.178 (P < 0.05), indicating deviation from Hardy-Weinberg equilibrium in these populations (Table 1).
AMOVA estimated F ST of 0.180 (P = 0.000) and R ST of 0.120 (P = 0.000), indicating moderate genetic divergence among populations (Table 2). According to pairwise comparisons, AFI followed by ESM population were the most divergent (Supplementary Table 1). The best number of clusters estimated by Bayesian clustering (Structure), considering both ΔK and mean likelihood methods, was K = 4 with each population been represented mostly by one genetic ancestry and varying admixture levels among them (Supplementary Fig. 1; Fig. 2). The CON followed by SER population showed higher admixture levels in its individuals (68% and 65% of individuals with Q < 0.9 for its most representative cluster, respectively), and AFI the lowest levels of admixture in the relation to other populations (Fig. 2). In addition, the Bayesian clustering showed that SER and CON were more related with each other, followed by ESM. AFI was the most divergent from other populations. These relationships are identified with K = 4 groups, but also when is considered grouping using K = 3, K = 5 and K = 6 genetic ancestries ( Supplementary Fig. 2). www.nature.com/scientificreports www.nature.com/scientificreports/ The contemporary N E estimated for the three populations (SER, CON, ESM) ranged from 13.5 to 22.7, indicating low number of reproductive individuals ( Table 3). The bottleneck test showed significant deviation of mutation-drift equilibrium in the three mutation models tested for all three populations (P < 0.05; Table 3), indicating that they suffered recent reductions in population size. The best historical migration model among the three populations was the full matrix migration (Supplementary Table 2). The values of the effective number of migrants (N E m) ranged from 0.086 from SER to ESM to 5.653 from ESM to SER (Supplementary Table 3). The estimated historical N E were 481, 760, and 11 for SER, CON, and ESM, respectively (Supplementary Table 3). species distribution models. Of the 19 bioclimatic variables from the WorldClim 29 , the following variables were retained to performing species distribution models after the factor analyses: mean diurnal range (Bio2), isothermality (Bio3), mean temperature of warmest quarter (Bio10), precipitation of wettest quarter (Bio16), and precipitation of driest quarter (Bio17). The values of TSS and AUC for model evaluation ranged from 0.333 to 1.000 and 0.611 to 1.000, respectively (Supplementary Tables 4 and 5). The variables that most contributed to the models were Bio3, Bio10, and Bio17 (Supplementary Table 6). The SDM projections confirmed the pattern of occurrence for D. exaltata, showing higher suitability in the semi-deciduous Atlantic Forest and in the Atlantic Forest/Cerrado transitional zone in southeastern Brazil, mainly in Minas Gerais, with smaller areas in São Paulo and Rio de Janeiro states (Fig. 3). The projections of both models for 2050 showed a loss in suitable area, mainly in the northern and northeastern parts of the range, which was more accentuated in the less-conservative BCC-CSM1-1 model (Fig. 3). In the most conservative scenario of climate change, MIROC-ESM, we observed a displacement of suitability toward the south, the opposite to a decrease in suitability in the southern area in the less-conservative model. There was also a decrease in suitability in the Atlantic Forest/Cerrado transitional zone, the area of most of the registered occurrences of the species (Fig. 3).
Geospatial conservation assessment. Using the Geospatial Conservation Assessment Tool software 30 (GeoCAT) based on 19 herbarium records to evaluating the conservation status of D. exaltata, we estimated an extent of occurrence (EOO) of 263 905 km 2 and a much smaller area of occupancy (AOO) of 76 km 2 for the species. Due to the possibility of misidentification in the only record for D. exaltata in speciesLink at Jequié, Bahia state, and because we could not find any other description for the species in Bahia in specialized literature 21,31,32 , we also performed the EOO and AOO analyses without this record. The estimates of EOO and AOO without Jequié were 174 936 km 2 and 72 km 2 , respectively, similar to estimates with the complete dataset. Based on the B criteria of the IUCN, the species was classified as Low Concern according to the EOO (EOO > 20 000 km 2 ) and    Table 3.  www.nature.com/scientificreports www.nature.com/scientificreports/ as Endangered according to the AOO (AOO < 500 km 2 ) in both analyses, with and without Jequié. In addition, following the B2 criteria, the species should also be considered Endangered because it meets the two following conditions: (a) severely fragmented populations and (b) continuous decline in area as indicated by (bi) projected EOO and (biii) continuing decline in area, extent, and/or quality of habitat.

Discussion
Our study showed that D. exaltata exhibits low genetic diversity, inbreeding, and evidence of a recent decrease in population size. Furthermore, species distribution models project a reduction in range size for the near future (2050). According to the geospatial criteria of the IUCN (2014), the species falls into the Endangered category of extinction risk. Altogether, our data indicate clearly that D. exaltata is a threatened species of the Atlantic Forest. The contribution of genetic data, in addition to ecological data, to establishing the extinction risk of the species and the importance of obtaining both types of data to establish measures of conservation are discussed below.
Dimorphandra exaltata populations have lower genetic diversity (H E = 0.289-0.560) than populations of several other legume tree species from the Atlantic Forest, including Hymenaea courbaril (H E = 0.536-0.804) 33 , [36][37][38] , and the vulnerable Dalbergia nigra (H E = 0.682-0.798) [39][40][41] , all species evaluated with microsatellite markers. Another appreciation of the level of genetic diversity of D. exaltata could be based on a comparison with closely related species, which discounts the phylogenetic effect 42,43 . Dimorphandra exaltata populations exhibited considerably lower genetic diversity (mean H E per population of 0.413) than a widespread congeneric species, D. mollis, which shows a mean H E per population of 0.594 44 , as expected due to its more restricted distribution. Dimorphandra exaltata populations showed even lower diversity than the narrowly endemic and critically endangered congeneric species D. wilsonii, with a mean H E per population of 0.559 27 . Although our genetic diversity evaluation was limited to 62 individuals, from three populations and isolated individuals from Atlantic Forest due to the rarity of the species, the low genetic diversity found indicates that D. exaltata is genetically depleted.
The contemporary N E of D. exaltata populations ranged from 13.5 to 22.7 and the estimated historical N E ranged from 11 to 700, which points to a low N E for the species. The comparison between the two N E together with the bottleneck tests point to a severe decrease in the population sizes of the species, what is probably associated with habitat loss and fragmentation in the study area. This pattern can explain the low diversity found in D. exaltata, since N E is one of the most relevant parameters to explain the patterns of genetic variation in wild populations 11 . These results are very important for the management of endangered species since N E of 100 and 1 000 individuals are suggested to prevent loss of genetic diversity by genetic drift for five generations and to maintain long-term evolutionary potential, respectively 45 .
The population genetic structure of D. exaltata suggests considerable admixture among populations, except for the isolated individuals of the Atlantic Forest. The historical demographic analysis reinforces that the three populations were genetically connected in the past. However, we should take care in extrapolating this connectivity to the current time. In fact, during our fieldwork, we observed a very low number of individuals and they were isolated and separated by large spatial distances, mainly in eastern Atlantic Forest. Also, the land use maps showed great anthropic pressure in the areas of occurrence of the species 46 (Fig. 1), which results in fragmented habitats and isolated populations. As consequence of this fragmentation, the gene flow of D. exaltata is likely compromised by a low-quality matrix among fragments. The seed dispersion of the species can be affected by a shortage of dispersers. Two species of the genus, D. mollis and D. wilsonii, exhibit low seed dispersal, which is mainly performed by tapirs (Tapirus terrestris), which are vulnerable and locally extinct in some areas 26 . Although the seed disperser of D. exaltata is not known, the similarity among species of the genus suggests a similar pattern for it. This is a matter of concern because at least 1.0 migrant per generation is necessary to counteract the genetic drift effects in the loss of diversity of populations 47 .
Our populations showed significant inbreeding coefficients, indicating a departure from random mating within populations. Although the mating system of D. exaltata is not known, it is probably similar to that of the congeneric D. wilsonii, which has self-compatibility 26 but mainly performs outcrossing 48 . The inbreeding found in D. exaltata may be due to selfing or mating between related individuals (biparental inbreeding). The small population size could favor mating among relatives or selfing in opposition to outcrossing. Increased selfing in preferentially outcrossing species is commonly found after fragmentation 5 . Furthermore, perennial plants, that have recently undergone habitat loss and fragmentation, such as D. exaltata, have a high probability of suffering inbreeding depression 8 . Therefore, inbreeding might be an additional threat to D. exaltata.
It is well-established that species with isolated and small populations are subject to loss of genetic diversity, a higher degree of inbreeding, and an elevated extinction risk in the wild 45 . Thus, the genetic data gathered here for D. exaltata, which showed low genetic diversity, inbreeding, and a recent reduction in population size, indicate that the "genetic health" of the species is compromised, which may decrease its adaptive capacity and threaten its survival in the wild in the near future. Thus, the genetic analyses of the species itself allow us to classify D. exaltata as a threatened species. In addition, the genetic data suggest that the species is more threatened than indicated by geospatial analyses.
The projected distribution for D. exaltata in the near future (2050) suggests a decrease in the total area of occurrence, including in the area in which it is most found currently, along with a displacement toward the south. The climate is considered to be the main driver of the species' range extent at large scales and of its long-term dynamics 44,[49][50][51][52][53] . In face of climate change, the probability of survival of a species is dependent on its capacity to adapt to the new environmental conditions or to migrate to suitable habitats 54 . However, the genetic data showed that the adaptive potential of D. exaltata is probably compromised. Thus, the species may depend on its migration capacity to colonize new favorable areas to survive in the face of climate change. However, the current www.nature.com/scientificreports www.nature.com/scientificreports/ connectivity among populations of the species is likely compromised by both low seed dispersal and pollen flow. Thus, the projected reduction of areas of occurrence and their displacement may be a concern for D. exaltata.
The geospatial assessment analyses showed that D. exaltata should be classified as Endangered, according the B2 criteria 4 (AOO less than 500 km 2 ). Furthermore, it could be also considered to be in the Threatened category because the populations are severely fragmented and there is a projected decline of this area, along with the low quality of its habitat (the sub criteria of B2). The Atlantic Forest has a fragmented distribution, with a history of disturbance and low functional connectivity among fragments 24 . Furthermore, the D. exaltata areas of occurrence in the Atlantic Forest/Cerrado transition zone, where the species is partially sympatric with D. wilsonii, are dominated by anthropogenic landscapes due to conversion of forested areas to agricultural lands, urbanization, and fire. These are the main threat factors for D. wilsonii 27 and may also play this role for D. exaltata.
Although D. exaltata occurs in one of the most-inventoried regions of southeastern Brazil, only 19 occurrence records were found, demonstrating the rarity of the species. In several areas of Atlantic Forest/Cerrado transition, D. exaltata occurs together with D. wilsonii, a critically endangered species, which has a National Action Plan for its conservation 27 (PAN faveiro de Wilson). It is highly recommended that this Action Plan also include D. exaltata, as the two species share similar ecological requirements and the same threat factors. Furthermore, only three records for D. exaltata were within protected areas. Protecting areas where the two Dimorphandra species co-occur will lead to protection of both threatened trees.
The genetic analyses indicated that the main risk factors to wild persistence of D. exaltata are its low genetic diversity and its low effective population size. Considering the ecological and genetic risk factors, conservation actions are urgently needed to decrease the probability of species extinction. Firstly, we recommend more search to find new individuals/populations of the species, mainly in easternmost areas Atlantic Forest. In addition, we recommend reintroduction, ex situ conservation, and genetic rescue to increase genetic diversity and maintain genetic connectivity among its populations. Genetic rescue is a highly beneficial strategy for management of genetically depleted populations 55 . For this, the creation and permanent maintenance of ex situ production of saplings from seeds of trees from different populations is a priority. As the analysed populations are not very genetically divergent, saplings from different populations could be introduced into the same locality, which could increase the genetic diversity and decrease the rate of inbreeding. However, considering the high genetic divergence of individuals from the eastern Atlantic Forest in relation to other areas, we suggested reintroduction in the eastern Atlantic Forest preferentially with saplings produced by seeds from these locations. The saplings should be used to increase the sizes of current populations, which must be protected, and to reintroduce the species into suitable areas according to SDM projections. It should be noted that the current suitable areas will suffer significant reduction in even the most optimistic projection of climate changes. Thus, it is recommended to take in account this projection when choosing localities for reintroduction.

Conclusion
Our integrative genetic and ecological framework to assess the degree to which a poorly known tree species is threatened and to guide conservation measures for that tree species, which occurs in the Atlantic Forest, a large and highly threatened biome in eastern South America, was demonstrated to be highly effective. This strategy of investigation, which includes genetic data, could be used for species in other conservation hotspots worldwide, where generally there is a lack of demographic information to establish the conservation status of species. From the results, our integrative framework supplies essential information for conservation policies.

Materials and Methods
study species. Dimorphandra exaltata is a medium to tall tree (15-30 meters), with bipinnate leaves, opposite leaflets pairs, and densely hairy petioles. Its inflorescences have a corymbe-type panicle consisting of multiple spikes. The flowers are cream or yellow with a glabrous chalice. Its fruits are flat legumes, which are almost straight, woody, and dark, indehiscent with sub-cylindrical seeds 21 . It flowers during November-December and fruit maturation occurs in September-October 21 . The pollinator and seed disperser of D. exaltata are not known. However, it has been proposed 21 that the pollination is performed by insects because of the similarity of its inflorescences and flowers with those of the congeneric species D. vernicosa, which is pollinated by bees of the Apidae family.

Herbarium records and study area.
Our study initially evaluated the occurrence of D. exaltata through analyses of herbarium records. Georeferenced herbarium records of D. exaltata were obtained mainly from the speciesLink database 56 , but to check for duplicates and to add missing records, other databases were used as well 21,31,32 . Due to the similarities among several species of the genus, the records of D. exaltata were compared with those of other Dimorphandra species to avoid misidentification. We retained only the most recent records by locality to avoid bias due to differential sampling efforts in certain areas and aggregated surveys. The localities were defined as the exact coordinates of the sites available in the record or alternatively, when the coordinate of the site was not available, by the municipalities coordinates. After the filtering process, we obtained only 19 georeferenced records in the speciesLink database (Supplementary Table 7 and Fig. 1), mostly in the semi-deciduous Atlantic Forest and in the Atlantic Forest/Cerrado transition zone. In the transition zone, the occurrences were mainly in cropland or urbanized areas 46 (Fig. 1). In addition, most of the peripheral records at the southern and eastern edges of occurrence were from near highly urbanized areas 46 (Fig. 1). For species distribution models (SDM) and geospatial conservation assessment analyses, we used the 19 records.
We also made a great effort to collect D. exaltata on Minas Gerais and Rio de Janeiro states, which concentrate the highest number of herbarium records and the most potential distribution areas for the species (Figs 1 and 3). In our collect expeditions in the eastern of Minas Gerais and the Rio de Janeiro, we found only four juveniles in www.nature.com/scientificreports www.nature.com/scientificreports/ one area and three isolated adults in two distant sites with distances ranging from 31.5 to 95.7 km apart from each other evidencing that the species is rare.
For the genetic analyses, we sampled 55 individuals of D. exaltata from three populations and seven isolated individuals from the three sites in eastern Minas Gerais and Rio de Janeiro, totaling 62 individuals (Fig. 1, Table 1). Two populations are located in Atlantic Forest/Cerrado transition areas, Contagem (CON) and Esmeraldas (ESM), where the species occurs in higher density and the third population is located in Serro (SER) in the semi-deciduous Atlantic Forest (Fig. 1). The isolated individuals of the species were in two sites already recorded in the speciesLink database for the species, Rio de Janeiro (RJA) and Valença (VAL) and in a municipality that do not had register for D. exaltata occurence, Santa Bárbara do Monte Verde (SBM) in the semi-deciduous Atlantic Forest. These areas are highly fragmented and disturbed due to urban pressure and soil conversion to croplands 46 (Fig. 1). The seven isolated individuals from RJA, VAL and SBM were used as representatives of the eastern Atlantic Forest, and were analysed together (AFI group), although they probably do not represent samples of only one population. AFI group was used only for estimates of genetic diversity and evaluation of genetic structure of species through of the analysis of molecular variance (AMOVA) and by the Bayesian clustering method implemented in STRUCTURE software 57,58 . The further genetic investigation was performed using only the populations SER, CON and ESM.
DNA isolation and microsatellite genotyping. Frozen leaf or vascular tissue was used for DNA extraction according to the cetyltrimethylammonium bromide (CTAB) protocol 59 , slightly modified as suggested by 60 . Individuals were genotyped for 11 microsatellite loci, five isolated in D. mollis 61 (Dmo5, Dmo7, Dmo13, Dmo20, Dmo21) and six isolated in D. wilsonii 62 (Dw21, Dw28, Dw33, Dw105, Dw52 and Dw103) Amplifications of all markers were performed according to 61 . The amplicons were submitted to capillary electrophoresis in an ABI prism 3500xl automated sequencer with GeneScan TM ROX500TM as a standard fragment (Applied Biosystems, Foster City, CA, USA) and genotyped in GeneMapper software Version 5.0 (Applied Biosystems). Scoring errors due to the presence of null alleles, stuttering, or large allele dropout were evaluated in MicroChecker version 2.2.0.2 28 . Linkage disequilibrium for each pair of loci for each population was tested using a randomization-based test in FSTAT version 2.9.3.2 63 .
Genetic status: genetic diversity and structure, migration, and effective population size. The genetic diversity of D. exaltata populations was estimated by the following parameters: mean number of alleles per locus (A), expected and observed heterozygosity (H E and H O respectively) using ARLEQUIN 3.5 64 and the allelic richness (A R ) using the rarefaction method of 65 in FSTAT version 2.9.3.2 63 . The number of private alleles were investigated in GenALEx 6.503 66,67 . The inbreeding coefficient (F IS ) was calculated and departures from Hardy-Weinberg expectations were tested with exact tests in FSTAT version 2.9.3.2 63 , with significance assessed after sequential Bonferroni correction for multiple comparisons. The F IS was not estimated for the AFI group because they can represent samples of more than one population. Analyses of molecular variance (AMOVA) implemented in ARLEQUIN 3.5 64 were used to estimate the hierarchical genetic structure and the pairwise F ST and R ST among populations.
The number of genetic clusters was evaluated using a Bayesian clustering method implemented in STRUCTURE version 2.3.4 57,58 . Parameter sets assumed an admixture model with correlated allele frequencies among patches. Ten independent simulations were run with 100 000 chains as burn-in and 1 000 000 Markov chain Monte Carlo (MCMC) replicates after burn-in, testing for K = 1-8. To evaluate the best K number, the mean log likelihood of the model through all runs and ΔK statistics were used 68 . To visualize the individual ancestry proportion, the average of ten runs for K was calculated in CLUMPP version 1.1.2 69 and the bar plot membership coefficient of each individual was built in DISTRUCT 1.1 70 .
We estimated the contemporary effective populations sizes (N E ) using the relation N E = 0.5/Θ 71 ; Θ is the group molecular coancestry for the total number of sampled trees in each population 72 . The calculation of Θ was made using the equation (1) ; F i is the inbreeding coefficient of the i-th individual, θ ij is the pairwise kinship between the individual i-th and j-th 38 and n is the sample size. F i and θ ij were estimated in SPAGeDi 1.1 software 73 . We evaluated for recent reductions in population size using a test for heterozygosity excess implemented in BOTTLENECK version 1.2.02 74 . Populations that have suffered a recent size decrease exhibit reductions of allele numbers and heterozygosity. However, reduction in allele diversity is faster than that in heterozygosity, which leads to an excess of heterozygosity under Hardy-Weinberg equilibrium in comparison with the expectations of mutation-drift equilibrium 75 . Bayesian simulations were carried out using three mutation models for microsatellites, infinite allele models (IAM), stepwise mutation model (SMM), and two-phase mutation model (TPM), with 30% multi-step mutations to avoid biases due to wrong assumptions about mutation models. We ran 1 000 000 MCMC and used the Wilcoxon sign-rank test to analyse the significance of differences in heterozygosities.
To test for models of historical gene flow among populations, we used a coalescent model implemented in MIGRATE-N 3.6.11 13 . Mutation-scaled effective population size (Θ = 4Ν E μ) and migration rates (M = m/μ), where μ is the mutation rate, were estimated using the Bayesian inference method. We considered μ = 4.76 × 10 −3 mutations per locus per generation for the microsatellite loci 76 . The Brownian motion model for microsatellite data was defined as the mutation process of the markers and the prior distribution was set as uniform with Θ (10 −5 to 100) and M (0 to 1 000). We ran 3 iterations with 2 000 000 MCMC after a burn-in of 50 000 generations, with sampling every 100 generations using a static heating scheme with four chains, swamping at every 10 chains based on an exponential temperature scheme. The independent runs were summarized to estimate the parameter values. The following models were tested: panmictic model, full migration model, zero-migration, a model with CON and ESM as one population, and a model with SER and CON as one population. We obtained the effective www.nature.com/scientificreports www.nature.com/scientificreports/ numbers of immigrants (N E m) using the equation (2) M * Θ = 4N E m, with the number 4 representing a scalar due to ploidy or inheritance of diploid nuclear data. The historical effective population size was estimated through the expression (3) (Θ/4μ) = Ν E . species distribution models. To assess the current potential distribution of D. exaltata, species distribution models (SDM) were performed using 19 bioclimatic variables available in the WorldClim database, with resolution of ~1 km 29 . To obtain a subset of uncorrelated explanatory variables, an exploratory factor analysis (EFA) was used in the psych 1.7.5 package implemented in the R environment 77,78 . The algorithms Maxent, Generalized Linear Models (GLM), Generalized Boosted Models (GBM), Artificial Neural Network (ANN), Classification tree analysis (CTA), Surface Range Envelopes (SRE), Flexible Discriminant Analysis (FDA), Multiple Adaptive Regression Splines (MARS), and Random Forest (RF) were implemented in the Biomod2 3.3-7 package in the R environment 79 . To evaluate model performance, we randomly subset points for training (75%) and points to test the predictive power of the models (25%). Then, we used the receiver-operating characteristic (ROC) curve and its area under curve (AUC) and the true skill statistic (TSS) to select the best models. Consensus maps were built based on five independent runs of each algorithm using the TSS as a criterion to discard low-predictive-power models (TSS > 0.7). To evaluate the impacts of future climate change on the distribution of the species, we projected its potential distribution in 2050 using the same set of variables for MIROC-ESM and BCC-CSM1-1 projections in an arc resolution of 30 s representing an intermediary and a maximum representative concentration pathway for this period, respectively. Geospatial conservation assessment. The geospatial conservation status of D. exaltata was evaluated using the extent of occurrence (EOO) and area of occupancy (AOO), as estimated using the Geospatial Conservation Assessment Tool software 30 (GeoCAT). Using this framework, we evaluated the conservation status of D. exaltata based on the International Union for the Conservation of Nature (IUCN) criteria B (EOO and AOO) to identify the threat category of the species 4 . The estimation of EOO was based on the minimum convex hull that contains all sites of known occurrence 30 . The estimation of AOO was based on the sum of squared grids of known occurrence based on grids of 2.0 km 2 as suggested by IUCN guidelines 4,30 . The same dataset used in the SDM was used for geospatial analysis.

Data Availability
The datasets generated during and/or analysed during the current study will be available in the DRYAD repository.