Climatic stability in the Brazilian Cerrado: implications for biogeographical connections of South American savannas,species richness and conservation in a biodiversity hotspot

Aim To investigate the historical distribution of the Cerrado across Quaternary climatic fluctuations and to generate historical stability maps to test: (1) whether the ‘historical climate’ stability hypothesis explains squamate reptile richness in the Cerrado; and (2) the hypothesis of Pleistocene connections between savannas located north and south of Amazonia. Location The Cerrado, a savanna biome and a global biodiversity hotspot distributed mainly in central Brazil. Methods We generated occurrence datasets from 1000 presence points randomly selected from the entire distribution of the Cerrado, as determined by two spatial definitions. We modelled the potential Cerrado distribution by implementing a maximum‐entropy machine‐learning algorithm across four time projections: current, mid‐Holocene (6 ka), Last Glacial Maximum (LGM, 21 ka) and Last Interglacial (LIG, 120 ka). We generated historical stability maps (refugial areas) by overlapping presence/absence projections of all scenarios, and checked consistencies with qualitative comparisons with available fossil pollen records. We built a spatially explicit simultaneous autoregressive model to explore the relationship between current climate, climatic stability, and squamate species richness. Results Models predicted the LGM and LIG as the periods of narrowest and widest Cerrado distributions, respectively, and were largely corroborated by palynological evidence. We found evidence for two savanna corridors (eastern coastal during the LIG, and Andean during the LGM) and predicted a large refugial area in the north‐eastern Cerrado (Serra Geral de Goiás refugium). Variables related to climatic stability predicted squamate richness better than present climatic variables did. Main conclusions Our results indicate that Bolivian savannas should be included within the Cerrado range and that the Cerrado’s biogeographical counterparts are not Chaco and Caatinga but rather the disjunct savannas of the Guyana shield plateaus. Climatic stability is a good predictor of Cerrado squamate richness, and our stability maps could be used in future studies to test diversity patterns and genetic signatures of different taxonomic groups and as a higher‐order landscape biodiversity surrogate for conservation planning.     

Squamate richness in the Brazilian Cerrado and its environmental–climatic associations

We investigate patterns of species richness of squamates (lizards, snakes, and amphisbaenians) in the Brazilian Cerrado, identifying areas of particularly high richness, and testing predictions of large‐scale richness hypotheses by analysing the relationship between species richness and environmental climatic variables. We used point localities from museum collections to produce maps of the predicted distributions for 237 Cerrado squamate species, using niche‐modelling techniques. We superimposed distributions of all species on a composite map, depicting richness across the ecosystem. Then, we performed a multiple regression analysis using eigenvector‐based spatial filtering (Principal Coordinate of Neighbour Matrices) to assess environmental–climatic variables that are best predictors of species richness. We found that the environmental–climatic and spatial filters multiple regression model explained 78% of the variation in Cerrado squamate richness (r² = 0.78; F = 32.66; P < 0.01). Best predictors of species richness were: annual precipitation, precipitation seasonality, altitude, net primary productivity, and precipitation during the driest quarter. A model selection approach revealed that several mechanisms related to the different diversity hypothesis might work together to explain richness variation in the Cerrado. Areas of higher species richness in Cerrado were located mainly in the south‐west, north, extreme east, and scattered areas in the north‐west portions of the biome. Partitioning of energy among species, habitat differentiation, and tolerance to variable environments may be the primary ecological factors determining variation in squamate richness across the Cerrado. High richness areas in northern Cerrado, predicted by our models, are still poorly sampled, and biological surveys are warranted in that region. The south‐western region of the Cerrado exhibits high species richness and is also undergoing high levels of deforestation. Therefore, maintenance of existing reserves, establishment of ecological corridors among reserves, and creation of new reserves are urgently needed to ensure conservation of species in these areas.     

Factors determining the habitat of Drosera sessilifolia in the humid zone of the Brazilian Cerrado

Drosera sessilifolia, a tiny carnivorous plant whose adult size is around 3 cm, sometimes inhabits the humid zone known as the campo limpo umido in the Brazilian Cerrado region. D. sessilifolia rarely occurs in the swampy area along the stream in this region; however, its occurrence is expected to decline with agricultural development. In the present study, we examined the factors determining the habitat and growth of D. sessilifolia in the campo limpo umido as well as how these factors have been modified by agricultural developments. Our survey suggests that the growth of D. sessilifolia is greatly influenced by light in areas where sufficient soil moisture is available. The light environment at the edge of a track constructed for reasons of agricultural development is in fact more suitable for D. sessilifolia than that found in the areas formerly inhabited by it. However, soon after construction, the artificial habitat may be occupied by tall-growing plants. In the former habitat of D. sessilifolia, the amount of accumulated litter and the size of the submerged spot also influenced its growth. In order to conserve short-growing plants, like D. sessilifolia in the plant community, the accompanying tall-growing dominant grasses should be controlled by periodic human disturbance.     

The lizard assemblage from Seasonally Dry Tropical Forest enclaves in the Cerrado biome, Brazil, and its association with the Pleistocenic Arc

Aim  To determine if the distributions of lizard species from Seasonally Dry Tropical Forest (SDTF) enclaves within the Cerrado biome in central Brazil are associated with the Tropical Seasonal Forests Region, a recently proposed phytogeographic unit of South America, corroborating the existence of a Pleistocenic Arc of SDTFs.
Location  SDTF remnants in the Paranã River valley, municipality of São Domingos, Goiás, Brazil.
Methods  Lizards were extensively sampled using haphazard sampling, funnel traps, and pitfall traps with drift fences during four expeditions. The composition of the SDTF lizard assemblage was compared with those from other South American phytogeographic regions (Caatinga, Cerrado, Chaco, Llanos, and the dry forests of Colombia and Bolivia), based on the literature and our own unpublished data.
Results  The SDTF lizard assemblage contained 20 species, including 11 species with extensive distributions among the regions considered, seven species shared exclusively with Cerrado localities, a single species shared exclusively with other SDTFs, and one endemic species. The presence of Lygodactylus klugei (Smith, Martin & Swain, 1977), presumably endemic to the Pleistocenic Arc formed by the Tropical Seasonal Forests Region, considerably extends the known distribution of this species, suggesting historical connections between Caatinga and Cerrado SDTF enclaves.
Main conclusions  The composition of the lizard assemblage in Cerrado SDTF enclaves seems to corroborate the recent proposal that the SDTF should be recognized as a phytogeographic unit (or dominium). The presence of disjunct populations and endemic species highlights the urgency of considering the uniqueness of the Paranã River valley SDTFs and the importance of its conservation.     

Conservation biogeography of anurans in Brazilian Cerrado

The increasing rates of declines in anuran populations worldwide are creating demands for urgent strategies to maximize conservation efforts. This may be critical in regions for which few detailed data on diversity, abundance and distribution are available, such as in the Cerrado of Central Brazil. In this paper, we used a macroecological approach based on the extent of occurrence of 131 species of Anura (Amphibia) in the Cerrado region to design a regional network of potential areas that preserves all anuran species. The final network, obtained using a simulation annealing algorithm based on complementarity, has a total of 17 cells, widely distributed throughout the biome. Minimum costs solutions were obtained in respect to total human population size, soybean production and bovine density, because these are the factors associated with human occupation that historically are more likely to cause broad scale habitat losses. The macro-scale approach used here can provide overall guidelines for conservation and define the focus for more local and effective conservation efforts.     

Tick fauna from two locations in the Brazilian savannah

The Cerrado is Brazil’s tropical savannah, which is arguably under greater threat than the Amazon rainforest. The Cerrado Biome of tropical South America covers about 2 million km² and is considered a biodiversity hot spot which means that it is especially rich in endemic species and particularly threatened by human activities. The Cerrado is increasingly exposed to agricultural activities which enhance the likelihood of mixing parasites from rural, urban and wildlife areas. Information about ticks from the Cerrado biome is scarce. In this report tick species free-living, on domestic animals and on a few wild animals in two farms in the Cerrado biome (Nova Crixás and Araguapaz municipalities, Goiás State, Brazil) are described. Amblyomma cajennense was the first and Amblyomma parvum the second host-seeking tick species found. Only two other tick species were found free-living: one Amblyomma nodosum and three Amblyomma naponense nymphs. Cattle were infested with Boophilus microplus and A. cajennense. Buffalos were infested with B. microplus and A. parvum. Dogs were infested with A. cajennense, Amblyomma ovale, A. parvum and Rhipicephalus sanguineus ticks. Anocentor nitens, B. microplus, A. cajennense, and A. parvum were found on horses. Amblyomma auricularium were found attached to nine-banded armadillos and Amblyomma rotundatum to red-footed tortoise, cururu toads and a rattlesnake. The latter was also infested with an adult A. cajennense. No tick was found on a goat, a tropical rat snake and a yellow armadillo. Among the observations the infestation of several domestic animals with A. parvum seems be the main feature. It suggests that this species might become a pest. However, the life cycle of A. parvum in nature, as well as its disease vectoring capacity, are largely unknown. It would be important to determine if it is a species expanding its geographic range by adaptation to new hosts or if it has been maintained in high numbers at definite locations by specific and still undetermined conditions. A higher prevalence of A. cajennense in most Brazilian biomes, with the exception of rainforests, was already shown before. Thus this species is favored by deforestation and is an important research target as it is the most common vector associated with the Brazilian spotted fever.     

A hawkmoth crossroads? Species richness,seasonality and biogeographical affinities of Sphingidae in a Brazilian Cerrado

Aim The aim of the study was (1) to describe the biodiversity of the sphingid assemblage in a Cerrado area in the Triângulo Mineiro region, south‐east of Brazil; (2) to evaluate the seasonal variations in species composition; (3) to compare the faunistic relationships between the Cerrado biome and adjacent ecosystems; and (4) to analyse the biogeographical pattern of species distribution in the Neotropical region in a historical context. Location Panga Ecological Station (PES), 30 km south of the city of Uberlândia, and other areas of the Triângulo Mineiro region, Minas Gerais state, south‐eastern Brazil. Methods Moth richness and abundance were monitored monthly at the PES from August 2003 to July 2004, with additional collections at this locality in 2001/2002, 2005 and 2006. Complementary moth richness and abundance data were also collected in other areas of the Triângulo Mineiro region. All collections were made using light traps, and the hawkmoths were mounted and identified. Cluster analysis, rarefaction curves and estimators of total species richness were used to compare the Cerrado hawkmoth assemblage with assemblages derived from other surveys in the Neotropics. Results In total, 61 hawkmoth species were recorded for the study region and their occurrence was markedly seasonal. The hawkmoth assemblage in the study area presented the closest similarity with rain forest areas and with a tropical dry forest area in Central America. The area shared species with both rain forest and seasonally dry tropical forest (SDTF) ecosystems, including supposedly endemic species previously recorded only in the latter areas. Rarefaction curves and estimators of the total number of species showed species richness to be comparable with other highly diverse forest areas in the Neotropics, such as the Brazilian Amazon and Costa Rica. Main conclusions This short‐term study is the first systematic survey of hawkmoths in the Cerrado. It has recorded around 22% of the South American fauna and highlighted the high species richness of the region, which compares favourably with that in other rain forest ecosystems. The survey indicates high regional diversity, and has shown that the Cerrado harbours a hawkmoth fauna comprising both rain forest elements, probably distributed along humid gallery forest corridors, and SDTF elements, supporting the idea of a historical Pleistocene arc connecting the Caatinga domain and other seasonal dry forest areas across the Cerrado region.     

Climate change, biofuels and eco-social impacts in the Brazilian Amazon and Cerrado

While global technical progress is relatively linear, there is wide variation in its environmental and social impacts at the local level, with cycles of expansion and retraction or boom and bust, of long or short duration. Analysis of previous open-ended stages of extraction and agro commodities in the Amazon indicates a general gravitational trend for technical progress to increase productivity and permit transformation of increasingly generic forms of material or energy, rather than relying on the specific physical or chemical properties provided by nature. While increased demand favours frontier expansion in the periphery when there is no other alternative, technical progress ultimately favours spatial reconcentration of production in central countries. The agroenergy stage now beginning involves rapid frontier expansion and offers various environmental and economic opportunities, but also generates a series of negative ecosystemic and socio-economic impacts, which are both direct and indirect, for tropical regions. The Amazon and the Cerrado are particularly vulnerable. Interacting with climate change and land use, the upcoming stage of cellulosic energy could result in a collapse of the new frontier into vast degraded pasture. The present and future impacts can be mitigated through crafting of appropriate policies, not limited to the Amazon, stressing intensified and more sustainable use of areas already cleared, minimizing new clearing and consolidation of alternatives for sustainable use of natural resources by local communities. Coping with these scenarios requires knowledge of complex causal relationships.     

Bat diversity of a Cerrado habitat in central Brazil

The Cerrado is considered one of the 25 biodiversity hotspots in the world by conservation organizations. There are few studies on bat taxonomic groups for this biome. Herein we present a bat survey employing mist nets in the protected area of a private natural heritage reserve Reserva Particular do Patrimônio Natural Pousada das Araras, located in the west-central Brazil. We investigate the hypothesis that the Cerrado habitat complexity plays a role on the different structuring forces in bat ecological communities. Bats represent a diversity of trophic levels, and they occupy a wide range of available Cerrado habitats and microhabitats. The patterns and processes we discuss represent the factors influencing coexisting species of bats in different habitats and their implications for conservation. We captured 758 individuals of 25 species belonging to four families. Phyllostomidae was the dominant family, represented by 20 species (80%). The average recapture rate was 6.2%, and the species with most proportional recapture was A. caudifer. Greater species richness was observed among bats with predominantly insectivore habits, followed by frugivores. Glossophaga soricina was a dominant species, with about 30% of the captures. There was seasonal variation relating to the number of bats captured, with greater bat frequency occurring during the wet season, although some species occurred at higher rates during the dry season, such as C. perspicillata and S. lilium. Sanguivore bats were abundant, reflecting the availability of shelters and food supply (livestock) in the surrounding area. Compared to other surveyed areas, Pousada das Araras may be considered of high species diversity, supplanting the majority of known Cerrado studied areas. Apparently Cerrado favours the occurrence of insectivore species, with emphasis on the foliage-gleaning insectivores belong to the subfamily Phyllostominae. This study indicates that apparently the conservation of the Cerrado savanna fragments can support a considerable diversity of bat species.     

The use of habitat mosaics by terrestrial vertebrate fauna: implications for conservation and management

Many species of vertebrates require multiple habitats to obtain different resources at different stages of their life-cycles. Use of habitat mosaics takes place on a variety of spatial and temporal scales, from a daily requirement for adjacent habitats to seasonal use of geographically separated environments. Mosaics of habitats are also required in some species to allow ontogenetic habitat shifts, while in others each sex may have specific requirements that are met by different habitats. The extent and nature of animal movements are key (but generally poorly known) factors affecting the vulnerability of species to landscape change. The requirement by many species for multiple habitats suggests that their conservation will be most effective in a mosaic environment and that protection of certain high profile habitats alone, such as rainforest, will be insufficient to achieve conservation goals. Management regimes that result in homogenization of habitats should be avoided. Priority should be given to research that identifies the extent to which species can locate habitat mosaics, at different spatial scales and arrangements, in modified environments.     

Mammal survey at a ranch of the Brazilian Cerrado

The wildlife of the Brazilian Cerrado is threatened by large-scale habitat loss, in particular due to conversion to agricultural land. It is essential to study how the mammal fauna copes with the highly fragmented, human-influenced, non-protected landscape. The paper presents the results of a survey of the large to medium-sized mammals of a typical cattle ranch with a mixture of human-created and natural vegetation types. We recorded 18 species. Surprisingly, several species were found to still thrive in the area, however, many species are rare or have become extinct. We conclude the paper with comments relevant for the conservation of mammals in the Cerrado as a whole.     

Evolutionary relationships between morphology, performance and habitat openness in the lizard genus Niveoscincus (Scincidae: Lygosominae)

Correlations between an animal's morphology and ecological parameters such as habitat characteristics emphasize the intimate link between phenotype and the environment, but are often difficult to interpret because the functional consequences of morphological variation are frequently unknown. We provide one of the few studies relating limb morphology, functional capabilities, and habitat in reptiles. We tested the hypothesis that species occupying open microhabitats would possess relatively longer limbs and faster sprint speeds than those occurring in more closed microhabitats. A number of morphological characteristics relevant to locomotion were quantified, including the length of the bones of the fore- and hindlimbs and body size. A phylogenetic analysis was then used to examine the evolutionary relationships between morphology, locomotor performance and microhabitat openness in seven species of Niveoscincus and one species of the closely related genus Pseudemoia. A significant evolutionary relationship was established between sprinting ability, morphology, and the openness of the microhabitat occupied by a species. The phylogenetic analysis demonstrated an evolutionary trend in Niveoscincus of species occupying open microhabitats (e.g. N. greeni, N ocellatus) being large with long limbs and high sprinting ability, while those occupying closed microhabitats (e.g. N coventryi, P. entrecasteauxii) art smaller with short limbs and much slower maximum sprint speeds.     

Influence of aquatic macrophyte habitat complexity on invertebrate abundance and richness in tropical lagoons

1. Aquatic plants are a key component of spatial heterogeneity in a waterscape, contributing to habitat complexity and helping determine diversity at various spatial scales. Theoretically, the more complex a habitat, the higher the number of species present. 2. Few empirical data are available to test the hypothesis that complexity increases diversity in aquatic communities (e.g. Jeffries, 1993 ). Fractal dimension has become widely applied in ecology as a tool to quantify the degree of complexity at different scales. 3. We investigated the hypothesis that complexity in vegetated habitat in two tropical lagoons mediates littoral invertebrate number of taxa (S) and density (N). Aquatic macrophyte habitat complexity was defined using a fractal dimension and a gradient of natural plant complexities. We also considered plant area, plant identity and, only for S, invertebrate density as additional explanatory variables. 4. Our results indicate that habitat complexity provided by the different architectures of aquatic plants, significantly affects both S and total N. However, number of individuals (as a result of passive sampling) also helps to account for S and, together with plant identity and area, contributes to the determination of N. We suggest that measurements of structural complexity, measured through fractal geometry, should be included in studies aimed at explaining attributes of attached invertebrates at small (e.g. plant or leaf) scales.     

Autoregressive modelling of species richness in the Brazilian Cerrado.

Spatial autocorrelation is the lack of independence between pairs of observations at given distances within a geographical space, a phenomenon commonly found in ecological data. Taking into account spatial autocorrelation when evaluating problems in geographical ecology, including gradients in species richness, is important to describe both the spatial structure in data and to correct the bias in Type I errors of standard statistical analyses. However, to effectively solve these problems it is necessary to establish the best way to incorporate the spatial structure to be used in the models. In this paper, we applied autoregressive models based on different types of connections and distances between 181 cells covering the Cerrado region of Central Brazil to study the spatial variation in mammal and bird species richness across the biome. Spatial structure was stronger for birds than for mammals, with R(2) values ranging from 0.77 to 0.94 for mammals and from 0.77 to 0.97 for birds, for models based on different definitions of spatial structures. According to the Akaike Information Criterion (AIC), the best autoregressive model was obtained by using the rook connection. In general, these results furnish guidelines for future modelling of species richness patterns in relation to environmental predictors and other variables expressing human occupation in the biome.     

Rapid restoration of belowground structure and fauna of a seagrass habitat

Seagrass meadows are habitat for an abundance and diversity of animal life, and their continuing global loss has focused effort on their restoration. This restoration not only aims to re‐establish the structure of the seagrass, but also to restore its function, particularly as habitat. The success of seagrass restoration is typically measured by the recovery of aboveground structure, but this ignores the important role of the belowground component of seagrass ecosystems, which may not recover at the same rate, and is equally important for faunal communities. We quantify infaunal communities (abundance, richness, and composition) within expanding plots of restored seagrass, and relate their change to the recovery of belowground seagrass biomass and sediment properties. We found that infaunal abundance and composition converged on that found in natural seagrass within 2 years, while the overall infaunal richness was not affected by habitat. The carbon content of surface sediments also recovered within 2 years, although recovery of belowground biomass and sediment grain size took 4 to 6 years. These results suggest that the structure of recovering seagrass habitats may not need to attain that of natural meadows before they support infauna that is comparable to natural communities. This pace and effectiveness of recovery provides some optimism for future seagrass restoration.     

Geographic variation in 'costs of reproduction' in the scincid lizard Lampropholis guichenoti

1. A major focus of life-history research has been the analysis of reproductive effort (RE). However, while clearly defined in theory, RE has proved very difficult to measure. Consequently, researchers have looked for indices that estimate the components of RE. Uncritical use of indices of reproductive investment, such as measures of costs of reproduction, without verification of the underlying assumptions of the link between these traits, may lead to spurious conclusions. In this paper, the common assumption that the physical burden of the clutch impairs locomotor ability in gravid reptiles is examined.
2. Two neighbouring populations of Australian scincid lizards ( Lampropholis guichenoti) are similar in adult body sizes, body shapes and reproductive output (egg sizes, clutch sizes, relative clutch masses).
3. Despite morphological and reproductive similarities, the effects of pregnancy on maternal locomotor ability (running speeds, as measured in a laboratory raceway) differed dramatically between the two populations.
4. Lizards from the two populations ran at similar speeds when nongravid (i.e. after egg-laying), but pregnancy significantly reduced running speeds in one population and increased them in the other. Thus, superficial similarities in body size and reproductive output masked a strong divergence in the locomotor 'costs' of reproduction.
5. Caution is advised if using simple measures of reproductive output (e.g. relative clutch mass) as indices of reproductive effort, or to generalize results even among conspecific populations.     

红树林水生动物栖息地功能及其渔业价值

红树林生长于热带、亚热带海陆交界的生态敏感带,其根系为生活于潮间带高度异质环境下的生物提供了适宜生境:藻类、双壳类、甲壳类等大量附生于红树根部;红树的呼吸根、支柱根、树干、倒落的枝条和残骸等,与沉积物形成的松软基质为大量底栖动物提供栖息地,根部结构间的空隙成为虾类、蟹类和鱼类等游泳动物的优良避难所和索饵场。红树林凋落物以碎屑形式进入食物网,连同浮游植物、附生藻类和底栖微藻等,是红树林生态系统的碳循环重要组成部分,为红树林区水生动物提供了丰富的食物来源。可见,食物来源丰富、隐蔽性强、捕食压力低等特点使得红树林成为水生动物的理想栖息地,许多水生动物选择在其中度过部分或完整生活史。另外,红树林也是重要经济动物(鱼、虾、蟹类)的育苗场,为近岸鱼类种群的补充和渔业活动提供支持。为合理开发红树林区渔业和有效保护红树林湿地生态系统,从生境价值、凋落物在红树林生态系统食物网中的贡献等方面总结了红树林栖息地功能及其渔业价值,提出今后的研究方向应将红树林的栖息地功能从其它河口、近岸栖息地中分离出来,甄别不同栖息地间的动态关系及其对渔业的影响。