Mammals in South American drylands: faunal similarity and trophic structure

We compared the fauna of small mammals (less than 500 g body weight) among five major South American drylands (Atacama, Altiplano, Monte, Patagonia and Caatinga) and found considerable heterogeneity and distinctiveness in species richness and composition between these biomes. From a total of 89 recorded species, 76 of them are restricted to only one of these drylands. The highland desert, or Altiplano, is the biome with the highest number of species. Despite the marked differences in the composition of the mammalian fauna, the trophic structure shows a rather consistent pattern: herbivores are the most important trophic group in all drylands. This consistency seems to be more the result of phylogenetic inertia than of similar ecological processes. Our results are compared with recent studies on desert small mammals across continents.     

Dioecy in the Caatinga,a Brazilian tropical dry forest: typical reproductive traits of a low frequent sexual system

Dioecy is a rare sexual system among angiosperms whose occurrence and frequency depend on the kind and geographical location of the ecosystem. We evaluate the frequency of dioecy in the Caatinga, a special semi-arid vegetation type in Brazil. We compare the floral traits and other ecological characteristics usually related to dioecy between the Caatinga and other tropical ecosystems. Dioecious species of the Caatinga were compiled based on 24 published floristic lists including most of the Caatinga domain. These lists were compared with publications comprising data on sexual systems in different kinds of vegetation in the world. Sexual system confirmation and the database of the reproductive traits of the dioecious species were made by consulting herbaria, literature, and field observations. We registered 22 dioecious species occurring in the Caatinga. The overall dioecy frequency was 2.36 %, considering all Caatinga species, and 6.91 % within woody species. Most dioecious species exhibited traits related to this sexual system (e.g., woody habit, greenish inconspicuous flowers, generalistic pollination syndrome, and zoochory). The distribution of dioecy across the Caatinga plant community exhibited a very low level of phylogenetic signal (D = ?0.145), indicating a phylogenetic structure under Brownian evolution (p = 0.671). The frequency of dioecy in the Caatinga (2.36 %) proved lower than reported for angiosperms in general (ca. 6 %) and for other tropical ecosystems. This low frequency can be related to particular ecological characteristics of the Caatinga, such as low percentage of trees >3–4 m tall (13–18.7 %), low rainfall (50 % of the Caatinga vegetation with annual rainfall lower than 750 mm), and strong seasonality (6–9 months of dry season). Dioecious species of the Caatinga exhibited typical reproductive traits for this sexual system, similar to what has been reported for dioecious species of other tropical ecosystems in the world.     

Speciation with gene flow in whiptail lizards from a Neotropical xeric biome

Two main hypotheses have been proposed to explain the diversification of the Caatinga biota. The riverine barrier hypothesis (RBH) claims that the São Francisco River (SFR) is a major biogeographic barrier to gene flow. The Pleistocene climatic fluctuation hypothesis (PCH) states that gene flow, geographic genetic structure and demographic signatures on endemic Caatinga taxa were influenced by Quaternary climate fluctuation cycles. Herein, we analyse genetic diversity and structure, phylogeographic history, and diversification of a widespread Caatinga lizard (Cnemidophorus ocellifer) based on large geographical sampling for multiple loci to test the predictions derived from the RBH and PCH. We inferred two well‐delimited lineages (Northeast and Southwest) that have diverged along the Cerrado–Caatinga border during the Mid‐Late Miocene (6–14 Ma) despite the presence of gene flow. We reject both major hypotheses proposed to explain diversification in the Caatinga. Surprisingly, our results revealed a striking complex diversification pattern where the Northeast lineage originated as a founder effect from a few individuals located along the edge of the Southwest lineage that eventually expanded throughout the Caatinga. The Southwest lineage is more diverse, older and associated with the Cerrado–Caatinga boundaries. Finally, we suggest that C. ocellifer from the Caatinga is composed of two distinct species. Our data support speciation in the presence of gene flow and highlight the role of environmental gradients in the diversification process.     

Primate Group Size and Abundance in the Caatinga Dry Forest, Northeastern Brazil

The Caatinga dry forest poses a series of ecological challenges for mammals in general and primates in particular. The erratic rainfall pattern impacts on plant diversity and phenological patterns; from year to year there is marked variability in fruit production and failure to fruit is common. The harshness apparently accounts for the impoverished mammalian fauna. However, data on primate abundance, distribution, and possible environmental effects on primate density are lacking in this type of dry forest. I censused the primate community in 3 habitats of the Serra da Capivara National Park, Piaui, NE Brazil, over a total distance of 318 km. Overall, the abundance of primates in the Caatinga dry forest is very low as a consequence of low abundance of food resources both in space and time. Alouatta caraya (predominantly folivorous) occurs at extremely low density, and during the dry season are apparently confined to canyon areas, where trees retain their leaves. Callithrix jacchus has morphological feeding specializations for gum-eating, and gum is an important resource during food bottleneck periods. Nonetheless, Callithrix jacchus occurs at comparatively low densities. Group sizes for howlers and marmosets in the Caatinga are significantly smaller than in other forest types. Contrarily, Cebus apella libidinosus had an average group size within the range reported for Amazonian and Atlantic forests. Researchers consider the generalized diet of capuchins as the explanation for their similar abundance in different habitats, indicating relative independence from ecological constraints. However, I suggest that capuchin foraging style and cognitive abilities are important factors accounting for their unreduced group size and density even under extreme conditions.     

High diversity of arbuscular mycorrhizal fungi in natural and anthropized sites of a Brazilian tropical dry forest (Caatinga)

Arbuscular mycorrhizal fungi (AMF) establish symbiotic associations with higher plants, which support the establishment and maintenance of plant communities across a range of environments, including those adversely affected by anthropogenic activity as well as natural sites. This study aimed at determining the diversity and distribution of AMF in areas of the tropical semi-arid region of Caatinga, Brazil, and compare areas in a naturally preserved state with anthropized sites. We characterized AMF communities in soil samples (n = 108), based on morphological taxonomy, at three sampling occasions and from six areas (typical Caatinga, extremely sandy Caatinga, stony Caatinga, rocky Caatinga, and two typical Caatinga areas that had been modified by human activities), at the National Park of Catimbau, Northeast Brazil. Eighty AMF species were recorded, with Glomus and Acaulospora predominating at all sites. There were significant differences in the composition of AMF communities between natural and anthropized sites, and among sampling occasions. Habitat-types also influenced AMF communities in Caatinga. Extensive tropical dry forest areas, such as the Catimbau National Park possess distinct niches, which maintain diverse AMF communities that are determined by anthropogenic activities, as well as vegetation types and environmental conditions.     

Nutrient dynamics within amazonian forest ecosystems

Summary A comparative analysis on the rate of fine litterfall and its associated nutrient fluxes was carried out in a mixed forest on Tierra Firme, a tall Amazon Caatinga and a Bana on podsolized sands near San Carlos de Rio Negro. There was seasonality in leaf fall and total litterfall in mixed forest and tall Amazon Caatinga forest but no definite trend in the Bana. Litterfall curves were significantly correlated among sites indicating common regulating factors in the three forests. Leaf litter from mixed forest on Tierra Firme was richer in N with extremely low Ca and Mg concentrations; tall Amazon Caatinga litter had higher P and Mg concentration, while Bana litter was low in N but K concentration was twice as high as in the other two forests. Annual fine litterfall in Tierra Firme mixed forest was nearly 4 times higher than in Bana, But N flux was 10 times higher, while Ca and Mg fluxes were similar. Tall Amazon Caatinga had Ca and Mg fluxes in litterfall 2–3 times higher than the other two forests. Within-stand efficiency of nitrogen, calcium and magnesium use, as measured by biomass/nutrient ratios, differentiates Tierra Firme from Caatinga and Bana forest: Tierra Firme has the lowest N, but the highest Ca and Mg use efficiencies. Higher P use efficiency was measured in Bana followed by Tierra Firme and Caatinga; while Tierra Firme and Caatinga showed similar higher K use efficiencies than Bana. N/P ratios indicates that Tierra Firme forest is limited by P availability, while low N availability predominates in Caatinga. Bana appears limited by both N and P. These differences probably relate to variations in degree of sclerophylly and leaf duration which determine leaf nutrient concentrations in the ecosystems studied.     

Are conservation units in the Caatinga biome,Brazil, efficient in the protection of biodiversity? An analysis based on the drosophilid fauna

Biodiversity is believed to be low in regions with vegetation that has adapted to water stress. Additionally, there is little interest by authorities in establishing and expanding conservation units in these areas. In Brazil, the Caatinga and the Cerrado biomes comprise this xerophilous vegetation. In both, climate is tropical and the dry season is long and well-defined. The Caatinga is the Brazilian biome with the smallest area protected by conservation units. This study evaluates the efficacy of conservation units in the Caatinga biome based on abundance and richness of drosophilids. Flies were collected inside and outside these areas. In total, approximately 23,000 flies of 32 species were collected in six conservation and six non-conservation sites. Two non-described species occurred exclusively inside protected areas, underlining the importance of conservation efforts in the maintenance of biodiversity. Other species were recorded exclusively outside conservation areas, which emphasizes the importance of establishing and expanding conservation units in the Caatinga. Native species were significantly more abundant inside conservation units, though the richness was similar in protected and non-protected areas. Abundance of exotic species outside conservation areas was statistically different in comparison with that of native ones.     

Human activities influence the occupancy probability of mammalian carnivores in the Brazilian Caatinga

The Caatinga is a semi‐arid domain, characterized by reduced humidity and high rates of anthropogenic impact. In addition to the low availability of water, carnivorous mammals are still exposed to a number of threats related to landscape modifications. We used data from camera traps and occupancy models to investigate the habitat use by carnivores in an area of Caatinga in northeastern Brazil. We found a negative correlation between the distance from wind farms and the occupancy probability of the jaguar, and a positive correlation with the occupancy probability of the jaguarundi. Puma and jaguarundi occupied primarily sites near watercourses, whereas the occupancy of the crab‐eating fox was correlated positively with the presence of poachers. The ocelot was detected more frequently at sites distant from human settlements, whereas the jaguar was detected more often in areas far from wind farms. We found a negative correlation between the distance of water and the detection of the ocelot. The detection of the crab‐eating fox was influenced positively by the detection of cattle. In addition to the negative influence of some anthropic activities, our results indicate that water is a very important resource for species, and the few permanent sources of this resource available in the area must be preserved. The replication of our research in other systems, worldwide, that are experiencing similar pressures, should permit a systematic evaluation of the management and conservation strategies needed to rebuild or maintain populations, restore ecosystems, and support conservation policies in human‐altered landscapes. Abstract in Portuguese is available with online material.     

Evidence for longitudinal migration by a “sedentary” Brazilian flycatcher,the Ash‐throated Casiornis

The digitalization of museum collections and concurrent increase in citizen‐science initiatives is ushering in an era of unprecedented availability of primary biodiversity data. These changes permit a reappraisal of phenological patterns of tropical species. I examined spatio‐temporal variation in the distribution patterns of an ostensibly sedentary endemic Brazilian flycatcher, the Ash‐throated Casiornis (Casiornis fuscus), using both specimen data from museums and sighting records and rich media data from citizen‐science initiatives. I found compelling evidence for partial intratropical longitudinal migration to Amazonia and the Cerrado biomes from the species’ core range in the semi‐desert Caatinga biome and adjacent ecotones. These records from outside of the Caatinga were distributed during the height of the dry season from April to October, although the Caatinga is not entirely vacated at this time. This pattern of partial migration leads to a doubling of the distributional range of Ash‐throated Casiornises and strongly suggests that the species is a breeding near‐endemic of the Caatinga biome. This pattern was potentially previously not apparent because of significant biases in specimen collection between biomes, giving a false sense of abundance in the Brazilian Amazon.     

Diversity and seasonal dynamic of a lizard assemblage in a Neotropical semiarid habitat

We assessed the lizard assemblage from a priority conservation Caatinga area from northeastern Brazil, through a pluriannual ecological approach, to expand the understanding on biodiversity patterns of Neotropical semiarid habitats. The studied area presented one of the richest lizard faunas among Caatinga sites, being composed primarily by species typical from open landscapes. The local species composition was more similar to assemblages from adjacent Caatinga ecoregions than to those from other areas within the same ecoregion. The inventoried lizard assemblage consisted of a few common species and a majority of low abundance ones, and its overall richness and abundance did not differ between rainy and dry months. Our findings demonstrated that the composition of lizard assemblages did not match with the current proposed Caatinga ecoregions, and revealed that the studied assemblage followed a lognormal species-abundance distribution, showing no significant seasonal fluctuation in richness and abundance.     

Biogeographical relationships among tropical forests in north-eastern Brazil

Aim To use parsimony analysis of endemicity and cladistic analysis of distributions and endemism to evaluate two hypotheses addressing biogeographical relationships among Amazonia, the Caatinga forest enclaves, Pernambuco Centre and the southern Atlantic Forest. Location North‐eastern Brazil, South America. Methods To find the most parsimonious areagram we analysed a matrix composed of the presence (1) or absence (0) of 745 taxa (i.e. 293 genera and 452 species of woody plants) within 16 localities belonging to the four large regions addressed in this study. Results One most parsimonious areagram was found and it shows a basal separation between the southern Atlantic Forest and all other regions. This break is followed by a separation between all Caatinga forest enclaves (except Baturité) from a cluster composed of Baturité, the Pernambuco Centre and Amazonia. In this cluster, the most basal separation isolates Baturité from the cluster formed by localities from Amazonia and the Pernambuco Centre. The biogeographical relationships among sites could not be explained by either a random distribution of species among sites or by the geographical distance between sites. Main conclusions We found strong cladistic signal within the raw distribution and phylogenetic data used in our analysis, indicating structured species assemblages in the surveyed localities. They have resulted from the fragmentation of an ancestral biota that was once widely distributed in the region. Our results also support the hypothesis that Atlantic Forest is not a biogeographically natural area, because the Pernambuco Centre is more closely related to Amazonia than to the southern Atlantic Forest. Finally, our data do not support the notion that Caatinga forest enclaves comprise a single biogeographical region, because one Caatinga forest enclave (Baturité) is much more closely related to the cluster formed by Amazonia and the Pernambuco Centre than to other sites. These relationships suggest the occurrence of forest connections between Amazonia and the Atlantic Forests across Caatinga during several periods of the Tertiary and Quaternary. However, palaeoecological data currently available for the Caatinga region are still scarce and do not have either the spatial or temporal resolution required to reconstruct the history of connections among the forests in north‐eastern Brazil.     

Plant sexual systems and a review of the breeding system studies in the Caatinga, a Brazilian tropical dry forest

BACKGROUNDS AND AIMS: The reproductive biology of a community can provide answers to questions related to the maintenance of the intraspecific pollen flow and reproductive success of populations, sharing and competition for pollinators and also questions on conservation of natural habitats affected by fragmentation processes. This work presents, for the first time, data on the occurrence and frequency of plant sexual systems for Caatinga communities, and a review of the breeding system studies of Caatinga species. METHODS: The sexual systems of 147 species from 34 families and 91 genera occurring in three Caatinga areas in north-eastern Brazil were analysed and compared with worldwide studies focusing on reproductive biology of different tropical communities. KEY RESULTS: The frequency of hermaphrodite species was 83.0 % (122 species), seven of these (or 4.8 % of the total) being heterostylous. Monoecy occurred in 9.5 % (14) of the species, and andromonoecy in 4.8 % (seven). Only 2.7 % (four) of the species were dioecious. A high percentage of hermaphrodite species was expected and has been reported for other tropical ecosystems. With respect to the breeding system studies with species of the Caatinga, the authors' data for 21 species and an additional 18 species studied by others (n = 39) revealed a high percentage (61.5 %) of obligatory self-incompatibility. Agamospermy was not recorded among the Caatinga studied species. CONCLUSIONS: The plant sexual systems in the Caatinga, despite the semi-arid climate, are similar to other tropical dry and wet forest communities, including those with high rainfall levels, except for the much lower percentage of dioecious species. The high frequency of self-incompatible species is similar to that reported for Savanna areas in Brazil, and also for dry (deciduous and semideciduous) and humid tropical forest communities.     

Manihot alterniflora and M. elongata spp. nov. (Euphorbiaceae) and the rediscovery of M. quinquefolia in Caatinga (semiarid) vegetation in Brazil

We describe and illustrate two species of Manihot that occur in Caatinga (semiarid) vegetation in Brazil and redescribe and lectotypify M. quinquefolia Pohl, which was only known from a single collection made by J. E. B. Pohl in 1827. Manihot elongata P.Carvalho & M.Martins is widely distributed and Manihot alterniflora P.Carvalho & M.Martins is endangered because of its small populations and restricted area of occurrence. We establish M. quinquefolia as the only species of Manihot in the Caatinga with compound leaves. An identification key is provided for the 13 species of Manihot present in the Caatinga.     

Identity and relationships of the Arboreal Caatinga among other floristic units of seasonally dry tropical forests (SDTFs) of north-eastern and Central Brazil

The tree species composition of seasonally dry tropical forests (SDTF) in north-eastern and central Brazil is analyzed to address the following hypotheses: (1) variations in species composition are related to both environment (climate and substrate) and spatial proximity; (2) SDTF floristic units may be recognized based on peculiar composition and environment; and (3) the Arboreal Caatinga, a deciduous forest occurring along the hinterland borders of the Caatinga Domain, is one of these units and its flora is more strongly related to the caatinga vegetation than to outlying forests. The study region is framed by the Brazilian coastline, 50th meridian west and 21st parallel south, including the Caatinga Domain and extensions into the Atlantic Forest and Cerrado Domains. Multivariate and geostatistic analyses were performed on a database containing 16,226 occurrence records of 1332 tree species in 187 georeferenced SDTF areas and respective environmental variables. Tree species composition varied significantly with both environmental variables and spatial proximity. Eight SDTF floristic units were recognized in the region, including the Arboreal Caatinga. In terms of species composition, its tree flora showed a stronger link with that of the Cerrado Dry Forest Enclaves. On the other hand, in terms of species frequency across sample areas, the links were stronger with two other units: Rock Outcrops Caatinga and Agreste and Brejo Dry Forests. There is a role for niche-based control of tree species composition across the SDTFs of the region determined primarily by the availability of ground water across time and secondarily by the amount of soil mineral nutrients. Spatial proximity also contributes significantly to the floristic cohesion of SDTF units suggesting a highly dispersal-limited tree flora. These units should be given the status of eco-regions to help driving the conservation policy regarding the protection of their biodiversity.     

The Role of Edaphic Environment and Climate in Structuring Phylogenetic Pattern in Seasonally Dry Tropical Plant Communities

Seasonally dry tropical plant formations (SDTF) are likely to exhibit phylogenetic clustering owing to niche conservatism driven by a strong environmental filter (water stress), but heterogeneous edaphic environments and life histories may result in heterogeneity in degree of phylogenetic clustering. We investigated phylogenetic patterns across ecological gradients related to water availability (edaphic environment and climate) in the Caatinga, a SDTF in Brazil. Caatinga is characterized by semiarid climate and three distinct edaphic environments – sedimentary, crystalline, and inselberg –representing a decreasing gradient in soil water availability. We used two measures of phylogenetic diversity: Net Relatedness Index based on the entire phylogeny among species present in a site, reflecting long-term diversification; and Nearest Taxon Index based on the tips of the phylogeny, reflecting more recent diversification. We also evaluated woody species in contrast to herbaceous species. The main climatic variable influencing phylogenetic pattern was precipitation in the driest quarter, particularly for herbaceous species, suggesting that environmental filtering related to minimal periods of precipitation is an important driver of Caatinga biodiversity, as one might expect for a SDTF. Woody species tended to show phylogenetic clustering whereas herbaceous species tended towards phylogenetic overdispersion. We also found phylogenetic clustering in two edaphic environments (sedimentary and crystalline) in contrast to phylogenetic overdispersion in the third (inselberg). We conclude that while niche conservatism is evident in phylogenetic clustering in the Caatinga, this is not a universal pattern likely due to heterogeneity in the degree of realized environmental filtering across edaphic environments. Thus, SDTF, in spite of a strong shared environmental filter, are potentially heterogeneous in phylogenetic structuring. Our results support the need for scientifically informed conservation strategies in the Caatinga and other SDTF regions that have not previously been prioritized for conservation in order to take into account this heterogeneity.     

Protected areas buffer the Brazilian semi‐arid biome from climate change

The Caatinga is a botanically unique semi‐arid ecosystem in northeast Brazil whose vegetation is adapted to the periodic droughts that characterize this region. However, recent extreme droughts events caused by anthropogenic climate change have challenged its ecological resilience. Here, we evaluate how deforestation and protection status affect the response of the Caatinga vegetation to drought. Specifically, we compared vegetation responses to drought in natural and deforested areas as well as inside and outside protected areas, using a time‐series of satellite‐derived Normalized Difference Vegetation Index (NDVI) and climatic data for 2008–2013. We observed a strong effect of deforestation and land protection on overall vegetation productivity and in productivity dynamics in response to precipitation. Overall, deforested areas had significantly lower NDVI and delayed greening in response to precipitation. By contrast, strictly protected areas had higher productivity and considerable resilience to low levels of precipitation, when compared to sustainable use or unprotected areas. These results highlight the importance of protected areas in protecting ecosystem processes and native vegetation in the Caatinga against the negative effects of climate change and deforestation. Given the extremely small area of the Caatinga currently under strict protection, the creation of new conservation areas must be a priority to ensure the sustainability of ecological processes and to avoid further desertification.     

Plant species from Brazilian Caatinga: a control alternative for Aedes aegypti

The mosquito Aedes aegypti is the main vector for the virus dengue, chikungunya and Zika. For its control, it is essential to search for natural products with insecticidal effects. The climatic singularity of Caatinga, an exclusive Brazilian biome, aids the survival of plants that produce secondary metabolites, which could be toxic to insects. Therefore, this review discusses the insecticidal potential of Caatinga plants on Aedes aegypti mosquitoes. The meta-analysis was performed using Review Manager Software 5.4.1®. Several studies have demonstrated the insecticidal efficacy of Caatinga plants on the egg, larvae, pupae and adult phases of Ae. aegypti, with a predominance of the plant activity in the larval stage. The leaves were the most utilized part of the plant. The essential oils from Caatinga plants were significantly active against Ae. aegypti (RR = 0.21, 95 % CI = 0.07 – 0.68, p = 0.009). The most promising botanical genera as an insecticide are: Abarema, Myracrodruon, Croton, Lippia and Syagrus. Among chemical compounds from these insecticidal plants has been identified and isolated flavonoids and fatty acids. Therefore, the Caatinga plant is a promising plant that contain bioactive compounds that are useful in the control of vector insects. This could contribute to the characterisation and valorisation of flora of this biome, as well as the production of environmentally friendly insecticides with specific action on target insects.     

Vegetation structure determines insect herbivore diversity in seasonally dry tropical forests

Vegetation structure can often determine insect herbivore fauna in forests, but this mechanism has been demonstrated in seasonally dry tropical forests (SDTFs) only at small spatial scales. In this study we evaluated the effects of the geographical location of SDTFs and vegetation structure on insect herbivore communities (leaf-chewing and sap-sucking guilds) in three Brazilian ecoregions (Cerrado, Cerrado/Caatinga transition, and Caatinga). We tested the following predictions: (1) insect herbivore species composition, richness, abundance and beta diversity differ among forests in different ecoregions; (2) insect richness, abundance and beta diversity are positively related to tree richness and density; (3) spatial turnover of species is the primary mechanism that generates herbivorous insect β-diversity in different ecoregions, and is positively influenced by tree richness. The composition, richness, and abundance of herbivorous insects differed over SDFs along the gradient of Cerrado and Caatinga. Both herbivore guilds responded positively to tree richness. Tree density only determined the richness and abundance of sap-sucking herbivores. Insect β-diversity was similar among Cerrado and transition areas, but lower in Caatinga itself; β-diversity was also positively affected by tree richness. Species turnover, as opposed to nestedness, was the main mechanism generating β-diversity, but itself was not related to tree richness. We demonstrate in this study the importance of landscape diversity and availability of local resources for herbivorous insect communities, and we emphasize the importance of SDTF conservation in different ecoregions as a result of species turnover.     

Phylogeographic structure of <Emphasis Type="Italic">Spondias tuberosa</Emphasis> Arruda Câmara (Anacardiaceae): seasonally dry tropical forest as a large and continuous refuge

Spondias tuberosa occurs in the Caatinga domain (seasonally dry tropical forest biome) of north-eastern Brazil, a large biome with ecogeographic regions that may have modelled the population structure of the species. Here we studied the phylogeographic pattern of S. tuberosa using sequences of the accD-psaI plastid region and six SSR markers in individuals distributed across 20 localities. The results for accD-psaI demonstrated nine haplotypes: some of which were exclusive to Caatinga ecoregions, whereas others were found in all localities. Spatial analysis of molecular variance revealed two groups (F_ct?=?0.34, P?<?0.0039) with 33.91% variation between them. The SSR analyses displayed 2–5 alleles at each locus, some of which were unique to certain localities. As in the accD-psaI region, the population structure obtained using SSR markers fell into two groups: (1) a large group containing the majority of the geographic region of Caatinga and (2) a small group near the Atlantic forest. We demonstrate the population structure of S. tuberosa, identifying the Caatinga as large, continuous refuge and the region near the interface between the Caatinga and the Atlantic forest as second refuge.