植物地理影响蝉类的生物地理分布

在位于美国德克萨斯州最西南的联合国教科文组织生物保护区内,研究了蝉类昆虫的分布格局与寄主植物分布的相关性。对该保护区(已被指定为美国国家公园)进行全面的蝉类标本采集,并精确记录采集地点、生境特征和植物种类等数据。将每种蝉的分布地与其各个采集地的植被类型建立相互对应的关系。该保护区共有蝉类昆虫6属12种(均为本地种)。其中,Tibicen chisosensis Davis是惟一的特有种,Diceroprocta canescens Davis和Okangodes terlingua Davis的分布稍超出保护区的边界。每种都与特定的植被类型紧密联系。蝉类昆虫的分布均限制在其寄主植物的分布范围内,据此,可推断出在特定的生境中会发现哪些蝉类物种。     

Patterns of morphology and behaviour in a cicada guild: A neutral model analysis

Morphological and behavioural patterns were examined in a guild of nine species of eastern Australian cicadas. Co-occurrence in, and structure of, 44 study plots were determined to relate morphological and behavioural patterns to habitat overlap and habitat structure. Multi-variate analyses were employed to generate relationships between the species. Guild substructure shown by the habitat analyses cannot be predicted from the group substructure indicated by the morphology/behaviour analyses. Therefore, extant combinations appear to be random collections from the nine-species pool insofar as morphology and behaviour are concerned. In particular, the most generalized species (in terms of habitat) are relatively extreme in morphology and behaviour, and vice versa. Despite these results, there are weak indications that at least two suites of characters exist, and that one of them is associated with each of the most extreme types of habitat. The reasons for these apparently contradictory results are discussed. There is no evidence to suggest that systematic morphological divergence has occurred in this guild.     

Environmental and neighbourhood effects on tree fern distributions in a neotropical lowland rain forest

Questions: To what extent are the distributions of tropical rain forest tree ferns (Cyatheaceae) related to environmental variation, and is habitat specialization likely to play a role in their local coexistence? Location: Lowland rain forest at La Selva Biological Station, Costa Rica. Methods: Generalized linear (GLM) and generalized additive (GAM) logistic regression were used to model the incidence of four tree fern species in relation to environmental and neighbourhood variables in 1154 inventory plots regularly distributed across 6 km² of old‐growth forest. Small and large size classes of the two most abundant species were modelled separately to see whether habitat associations change with ontogeny. Results: GLM and GAM model results were similar. All species had significant distributional biases with respect to micro‐habitat. Environmental variables describing soil variation were included in the models most often, followed by topographic and forest structural variables. The distributions of small individuals were more strongly related to environmental variation than those of larger individuals. Significant neighbourhood effects (spatial autocorrelation in intraspecific distributions and non‐random overlaps in the distributions of certain species pairs) were also identified. Overlaps between congeners did not differ from random, but there was a highly significant overlap in the distributions of the two most common species. Conclusions: Our results support the view that habitat specialization is an important determinant of where on the rain forest landscape tree ferns grow, especially for juvenile plants. However, other factors, such as dispersal limitation, may also contribute to their local coexistence.     

Metacommunity patterns in larval odonates

The growth of metacommunity ecology as a subdiscipline has increased interest in how processes at different spatial scales structure communities. However, there is still a significant knowledge gap with respect to relating the action of niche- and dispersal-assembly mechanisms to observed species distributions across gradients. Surveys of the larval dragonfly community (Odonata: Anisoptera) in 57 lakes and ponds in southeast Michigan were used to evaluate hypotheses about the processes regulating community structure in this system. We considered the roles of both niche- and dispersal-assembly processes in determining patterns of species richness and composition across a habitat gradient involving changes in the extent of habitat permanence, canopy cover, area, and top predator type. We compared observed richness patterns and species distributions in this system to patterns predicted by four general community models: species sorting related to adaptive trade-offs, a developmental constraints hypothesis, dispersal assembly, and a neutral community assemblage. Our results supported neither the developmental constraints nor the neutral-assemblage models. Observed patterns of richness and species distributions were consistent with patterns expected when adaptive tradeoffs and dispersal-assembly mechanisms affect community structure. Adaptive trade-offs appeared to be important in limiting the distributions of species which segregate across the habitat gradient. However, dispersal was important in shaping the distributions of species that utilize habitats with a broad range of hydroperiods and alternative top predator types. Our results also suggest that the relative importance of these mechanisms may change across this habitat gradient and that a metacommunity perspective which incorporates both niche- and dispersal-assembly processes is necessary to understand how communities are organized. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dynamics of periodical cicada chorus centers (Homoptera: Cicadidae:Magicicada)

Mating aggregations of three species of periodical cicadas were monitored during the emergence of Brood XIX at a 16-ha study site in northwest Arkansas, May–June 1985. Magicicada tredecassiniappeared first and formed the most choruses. M. tredecimand M. tredeculachoruses formed next, and M. tredeculachoruses outnumbered those of M. tredecim.Of the 268 choruses seen, 84% were composed of M. tredecassini. M. tredecassiniwere often found chorusing in the same trees with the other two species. Such multispecies mating aggregations apparently are unique to periodical cicadas. Choruses were dynamic with respect to their locations and durations. Initially, choruses were located near areas of high cicada emergence densities. One week later, cicadas chorused in trees throughout the forest and at the forest edge. Many choruses were seen only once at a location. Although cicadas chorused for almost 4 weeks, individual choruses persisted only approximately 8 days, on average. Sound intensities under chorus centers ranged from 50 to 80 dh and were correlated with arena sizes during times of peak chorus activity. No distinct habitat preferences of the three species were observed, however, the tree species used by chorusing cicadas differed among the species.     

Alpine bird distributions along elevation gradients: the consistency of climate and habitat effects across geographic regions

Many species have shown recent shifts in their distributions in response to climate change. Patterns in species occurrence or abundance along altitudinal gradients often serve as the basis for detecting such changes and assessing future sensitivity. Quantifying the distribution of species along altitudinal gradients acts as a fundamental basis for future studies on environmental change impacts, but in order for models of altitudinal distribution to have wide applicability, it is necessary to know the extent to which altitudinal trends in occurrence are consistent across geographically separated areas. This was assessed by fitting models of bird species occurrence across altitudinal gradients in relation to habitat and climate variables in two geographically separated alpine regions, Piedmont and Trentino. The ten species studied showed non-random altitudinal distributions which in most cases were consistent across regions in terms of pattern. Trends in relation to altitude and differences between regions could be explained mostly by habitat or a combination of habitat and climate variables. Variation partitioning showed that most variation explained by the models was attributable to habitat, or habitat and climate together, rather than climate alone or geographic region. The shape and position of the altitudinal distribution curve is important as it can be related to vulnerability where the available space is limited, i.e. where mountains are not of sufficient altitude for expansion. This study therefore suggests that incorporating habitat and climate variables should be sufficient to construct models with high transferability for many alpine species.     

Endothermy in African platypleurine cicadas: the influence of body size and habitat (Hemiptera: Cicadidae)

The platypleurine cicadas have a wide distribution across Africa and southern Asia. We investigate endothermy as a thermoregulatory strategy in 11 South African species from five genera, with comparisons to the lone ectothermic platypleurine we found, in an attempt to ascertain any influence that habitat and/or body size have on the expression of endothermy in the platypleurine cicadas. Field measurements of body temperature (T(b)) show that these animals regulate T(b) through endogenous heat production. Heat production in the laboratory elevated T(b) to the same range as in animals active in the field. Maximum T(b) measured during calling activity when there was no access to solar radiation ranged from 13.2 degrees to 22.3 degrees C above ambient temperature in the five species measured. The mean T(b) during activity without access to solar radiation did not differ from the mean T(b) during diurnal activity. All platypleurines exhibit a unique behavior for cicadas while warming endogenously, a temperature-dependent telescoping pulsation of the abdomen that probably functions in ventilation. Platypleurines generally call from trunks and branches within the canopy and appear to rely on endothermy even when the sun is available to elevate T(b), in contrast to the facultative endothermy exhibited by New World endothermic species. The two exceptions to this generalization we found within the platypleurines are Platypleura wahlbergi and Albanycada albigera, which were the smallest species studied. The small size of P. wahlbergi appears to have altered their thermoregulatory strategy to one of facultative endothermy, whereby they use the sun when it is available to facilitate increases in T(b). Albanycada albigera is the only ectothermic platypleurine we found. The habitat and host plant association of A. albigera appear to have influenced the choice of ectothermy as a thermoregulatory strategy, as the species possesses the metabolic machinery to elevate to the T(b) range observed in the endothermic species. Therefore, size and habitat appear to influence the expression of thermoregulatory strategies in African platypleurine cicadas.     

The contributions of topoclimate and land cover to species distributions and abundance: fine‐resolution tests for a mountain butterfly fauna

Aim Models relating species distributions to climate or habitat are widely used to predict the effects of global change on biodiversity. Most such approaches assume that climate governs coarse‐scale species ranges, whereas habitat limits fine‐scale distributions. We tested the influence of topoclimate and land cover on butterfly distributions and abundance in a mountain range, where climate may vary as markedly at a fine scale as land cover. Location Sierra de Guadarrama (Spain, southern Europe) Methods We sampled the butterfly fauna of 180 locations (89 in 2004, 91 in 2005) in a 10,800 km² region, and derived generalized linear models (GLMs) for species occurrence and abundance based on topoclimatic (elevation and insolation) or habitat (land cover, geology and hydrology) variables sampled at 100‐m resolution using GIS. Models for each year were tested against independent data from the alternate year, using the area under the receiver operating characteristic curve (AUC) (distribution) or Spearman's rank correlation coefficient (r_s) (abundance). Results In independent model tests, 74% of occurrence models achieved AUCs of > 0.7, and 85% of abundance models were significantly related to observed abundance. Topoclimatic models outperformed models based purely on land cover in 72% of occurrence models and 66% of abundance models. Including both types of variables often explained most variation in model calibration, but did not significantly improve model cross‐validation relative to topoclimatic models. Hierarchical partitioning analysis confirmed the overriding effect of topoclimatic factors on species distributions, with the exception of several species for which the importance of land cover was confirmed. Main conclusions Topoclimatic factors may dominate fine‐resolution species distributions in mountain ranges where climate conditions vary markedly over short distances and large areas of natural habitat remain. Climate change is likely to be a key driver of species distributions in such systems and could have important effects on biodiversity. However, continued habitat protection may be vital to facilitate range shifts in response to climate change.     

Geographic body size variation in the periodical cicadas Magicicada: implications for life cycle divergence and local adaptation

Seven species in three species groups (Decim, Cassini and Decula) of periodical cicadas (Magicicada) occupy a wide latitudinal range in the eastern United States. To clarify how adult body size, a key trait affecting fitness, varies geographically with climate conditions and life cycle, we analysed the relationships of population mean head width to geographic variables (latitude, longitude, altitude), habitat annual mean temperature (AMT), life cycle and species differences. Within species, body size was larger in females than males and decreased with increasing latitude (and decreasing habitat AMT), following the converse Bergmann's rule. For the pair of recently diverged 13‐ and 17‐year species in each group, 13‐year cicadas were equal in size or slightly smaller on average than their 17‐year counterparts despite their shorter developmental time. This fact suggests that, under the same climatic conditions, 17‐year cicadas have lowered growth rates compared to their 13‐years counterparts, allowing 13‐year cicadas with faster growth rates to achieve body sizes equivalent to those of their 17‐year counterparts at the same locations. However, in the Decim group, which includes two 13‐year species, the more southerly, anciently diverged 13‐year species (Magicicada tredecim) was characterized by a larger body size than the other, more northerly 13‐ and 17‐year species, suggesting that local adaptation in warmer habitats may ultimately lead to evolution of larger body sizes. Our results demonstrate how geographic clines in body size may be maintained in sister species possessing different life cycles.     

Comments on the morphology of Nicsmirnovius eximius (Kiser, 1948) (Aloninae,Anomopoda, Cladocera) in Thailand,with description of its male

The density, diet and habitat use of brown trout (Salmo trutta) and Siberian sculpin (Cottus poecilopus) were studied in the subalpine River Atna in southeastern Norway in the autumn during a six year period (1986–1991). There was an inverse relationship between the density of brown trout and Siberian sculpin. Diet overlap, as indicated by the Schoener index, was high between the two species, ranging between 0.48 and 0.86. Chironomid larvae and other aquatic insects were the most common food items for both species. Brown trout also consumed substantive amounts of surface insects. Siberian sculpin typically occupied sites with finer substrates and greater water depths than brown trout, even though there was considerable overlap in habitat use between the two species. Because the two species shared similar habitats, we suggest that the potential for species interactions exists, particularly at sites where density of sculpin is high.     

The relative importance of climate and habitat in determining the distributions of species at different spatial scales: a case study with ground beetles in Great Britain

Experimental studies have shown that many species show preferences for different climatic conditions, or may die in unsuitable conditions. Climate envelope models have been used frequently in recent years to predict the presence and absence of species at large spatial scales. However, many authors have postulated that the distributions of species at smaller spatial scales are determined by factors such as habitat availability and biotic interactions. Climatic effects are often assumed by modellers to be unimportant at fine resolutions, but few studies have actually tested this. We sampled the distributions of 20 beetle species of the family Carabidae across three study sites by pitfall trapping, and at the national scale from monitoring data. Statistical models were constructed to determine which of two sets of environmental variables (temperature or broad habitat type) best accounted for the observed data at the three sites and at the national (Great Britain) scale. High‐resolution temperature variables frequently produced better models (as determined by AIC) than habitat features when modelling the distributions of species at a local scale, within the three study sites. Conversely, habitat was always a better predictor than temperature when describing species’ distributions at a coarse scale within Great Britain. Northerly species were most likely to occur in cool micro‐sites within the study sites, whereas southerly species were most likely to occur in warm micro‐sites. Effects of microclimate were not limited to species at the edges of their distribution, and fine‐resolution temperature surfaces should therefore ideally be utilised when undertaking climate‐envelope modelling.     

Changes in potential habitat of 147 North American breeding bird species in response to redistribution of trees and climate following predicted climate change

Mounting evidence shows that organisms have already begun to respond to global climate change. Advances in our knowledge of how climate shapes species distributional patterns has helped us better understand the response of birds to climate change. However, the distribution of birds across the landscape is also driven by biotic and abiotic components, including habitat characteristics. We therefore developed statistical models of 147 bird species distributions in the eastern United States, using climate, elevation, and the distributions of 39 tree species to predict contemporary bird distributions. We used randomForest, a robust regression‐based decision tree ensemble method to predict contemporary bird distributions. These models were then projected onto three models of climate change under high and low emission scenarios for both climate and the projected change in suitable habitat for the 39 tree species. The resulting bird species models indicated that breeding habitat will decrease by at least 10% for 61–79 species (depending on model and emissions scenario) and increase by at least 10% for 38–52 species in the eastern United States. Alternatively, running the species models using only climate/elevation (omitting tree species), we found that the predictive power of these models was significantly reduced (p<0.001). When these climate/elevation‐only models were projected onto the climate change scenarios, the change in suitable habitat was more extreme in 60% of the species. In the end, the strong associations with vegetation tempers a climate/elevation‐only response to climate change and indicates that refugia of suitable habitat may persist for these bird species in the eastern US, even after the redistribution of tree species. These results suggest the importance of interacting biotic processes and that further fine‐scale research exploring how climate change may disrupt species specific requirements is needed.     

Vanishing islands in the sky? A comparison of correlation- and mechanism-based forecasts of range dynamics for montane salamanders under climate change

Forecasting the effects of climate change on species and populations is a fundamental goal of conservation biology, especially for montane endemics which seemingly are under the greatest threat of extinction given their association with cool, high elevation habitats. Species distribution models (also known as niche models) predict where on the landscape there is suitable habitat for a species of interest. Correlative niche modeling, the most commonly employed approach to predict species' distributions, relies on correlations between species' localities and current environmental data. This type of model could spuriously forecast less future suitable habitat because species' current distributions may not adequately represent their thermal tolerance, and future climate conditions may not be analogous to current conditions. We compared the predicted distributions for three montane species of Plethodon salamanders in the southern Appalachian Mountains of North America using a correlative modeling approach and a mechanistic model. The mechanistic model incorporates species-specific physiology, morphology and behavior to predict an annual energy budget on the landscape. Both modeling approaches performed well at predicting the species' current distributions and predicted that all species could persist in habitats at higher elevation through 2085. The mechanistic model predicted more future suitable habitat than the correlative model. We attribute these differences to the mechanistic approach being able to model shifts in key range-limiting biological processes (changes in surface activity time and energy costs) that the correlative approach cannot. Choice of global circulation model (GCM) contributed significantly to distribution predictions, with a tenfold difference in future suitability based on GCM, indicating that GCM variability should be either directly included in models of species distributions or, indirectly, through the use of multi-model ensemble averages. Our results indicate that correlative models are over-predicting habitat loss for montane species, suggesting a critical need to incorporate mechanisms into forecasts of species' range dynamics.     

Comparative thermal adaptation in cicadas (Hemiptera: Cicadidae) inhabiting Mediterranean ecosystems

The thermal responses of cicadas inhabiting the Mediterranean ecosystems in Europe, North America, South Africa, and Australia are investigated. A total of 37 species and two subspecies from 17 genera representing eight tribes and three subfamilies of cicadas are investigated. The analysis includes species that are restricted to the Mediterranean ecosystem as well as those which also inhabit additional environments. The data suggest that cicadas adapt to the climate type regardless of particular types of plants within the various Mediterranean communities. Similarly, cicada thermal responses are independent of body size or taxonomic affinities. There is a wider range of body temperatures for the maximum voluntary tolerance temperature than for heat torpor or minimum flight temperatures. This diversity seems to be determined by the subdivision of the habitat used and the behavior of the species. All species possess relatively elevated heat torpor temperatures adapting to the general thermal characteristics of the Mediterranean ecosystem. The data suggest that cicadas adapt to the Mediterranean climate type regardless of the diversity of particular types of plants within the various communities, of body size or of taxonomic position.     

Palaeodistribution modelling does not support disjunct Pleistocene refugia in several Central American plant taxa

Aim We combine evidence from palaeoniche modelling studies of several tree species to estimate the extent of Central American forest during the Last Glacial Maximum (LGM). In particular, we ask whether the distributions of these species are likely to have changed since the LGM, and whether LGM distributions coincide with previously proposed Pleistocene refugia in this area. Location Central American wet and seasonally dry forests. Methods We developed ecological niche models using two simulations of Pleistocene climate and occurrence data for 15 Neotropical plant species. We focused on palaeodistribution models of three ‘focal’ tree species that occur in wet and seasonally dry Central American forests, where recent phylogeographic data suggest Pleistocene differentiation coincident with previously proposed refugia. We added predictions from six wet‐forest and six seasonally dry‐forest obligate plant species to gauge whether Pleistocene range shifts were specific to habitat type. Correlation analyses were performed between projected LGM and present distributions, LGM distributions and previously proposed refugia. We also asked whether modelled palaeodistributions were smaller than their current extents. Results According to our models, the ranges of the study species were not reduced during the LGM, and did not correlate with refugial models, regardless of habitat type. Relative range sizes between present and LGM distributions did not indicate significant range changes since the LGM. However, relative range sizes differed overall between the two palaeoclimate models. Main conclusions Many of the modelled palaeodistributions of study species were not restricted to refugia during the LGM, regardless of forest type. While constrained from higher elevations, most species found suitable habitat at coastal margins and on newly exposed land due to lowered sea levels during the LGM. These results offer no corroboration for Pleistocene climate change as a driver of genetic differentiation in the ‘focal’ species. We offer alternative explanations for genetic differentiation found in plant species in this area.     

Presence-absence versus presence-only modelling methods for predicting bird habitat suitability

Habitat suitability models can be generated using methods requiring information on species presence or species presence and absence. Knowledge of the predictive performance of such methods becomes a critical issue to establish their optimal scope of application for mapping current species distributions under different constraints. Here, we use breeding bird atlas data in Catalonia as a working example and attempt to analyse the relative performance of two methods: the Ecological Niche factor Analysis (ENFA) using presence data only and Generalised Linear Models (GLM) using presence/absence data. Models were run on a set of forest species with similar habitat requirements, but with varying occurrence rates (prevalence) and niche positions (marginality). Our results support the idea that GLM predictions are more accurate than those obtained with ENFA. This was particularly true when species were using available habitats proportionally to their suitability, making absence data reliable and useful to enhance model calibration. Species marginality in niche space was also correlated to predictive accuracy, i.e. species with less restricted ecological requirements were modelled less accurately than species with more restricted requirements. This pattern was irrespective of the method employed. Models for wide‐ranging and tolerant species were more sensitive to absence data, suggesting that presence/absence methods may be particularly important for predicting distributions of this type of species. We conclude that modellers should consider that species ecological characteristics are critical in determining the accuracy of models and that it is difficult to predict generalist species distributions accurately and this is independent of the method used. Being based on distinct approaches regarding adjustment to data and data quality, habitat distribution modelling methods cover different application areas, making it difficult to identify one that should be universally applicable. Our results suggest however, that if absence data is available, methods using this information should be preferably used in most situations.     

Habitat specificity and diversity of tree species in an African wet tropical forest

Niche differentiation with respect to habitat has been hypothesized to shape patterns of diversity and species distributions in plant communities. African forests have been reported to be relatively less diverse compared to highly diversed regions of the Amazonian or Southeast Asian forests, and might be expected to have less niche differentiation. We examined patterns of structural and floristic differences among five topographically defined habitats for 494 species with stems ≥1 cm dbh in a 50-ha plot in Korup National Park, Cameroon. In addition, we tested for species–habitat associations for 272 species (with more than 50 individuals in the plot) using Torus translation randomization tests. Tree density and basal area were lowest in areas with negative convexity, which contained streams or were inundated during rainy periods and highest in moist well-drained habitats. Species composition and diversity varied along the topographical gradient from low flat to ridge top habitats. The low depression and low flat habitats were characterized by high diversity and similar species composition, relative to slopes, high gullies and ridge tops. Sixty-three percent of the species evaluated showed significant positive associations with at least one of the five habitat types. The majority of associations were with low depressions (75 species) and the fewest with ridge tops (8 species). The large number of species–habitat associations and the pronounced contrast between low (valley) and elevated (ridgetop) habitats in the Korup plot shows that niche differentiation with respect to edaphic variables (e.g., soil moisture, nutrients) contributes to local scale tree species distributions and to the maintenance of diversity in African forests.     

Species–habitat associations and demographic rates of forest trees

Niche‐driven effects on demographic processes generated in response to habitat heterogeneity partly shape local distributions of species. Thus, tree distributions are commonly studied in relation to habitat conditions to understand how niche differentiation contributes to species coexistence in forest communities. Many such studies implicitly assume that local abundance reflects habitat suitability, and that abundance is relatively stable over time. We compared models based on abundance with those based on demographic performance for making inferences about habitat association for 287 tree species from three large dynamic plots located in tropical, subtropical and temperate forests. The correlation between the predictions of the abundance‐based models and the demography‐based models varied widely, with correlation coefficients ranging nearly from ?1 to 1.This suggests that the two types of models capture different information about species–habitat associations. Demography‐based models evaluate habitat quality by focusing on population processes and thus should be preferred for understanding responses of tree species to habitat conditions, especially when habitat conditions are changing and species–habitat interactions cannot be considered to be at equilibrium.     

Evaluating models to assess the distribution of Buxus balearica in southern Spain

Question: Which is the best model to predict the habitat distribution of Buxus balearica Lam. in southern Spain? Location: Málaga and Granada, Spain, across an area of 38 180 km². Methods: Prediction models based on 17 environmental variables were tested. Six methods were compared: multivariate adaptive regression spline (MARS), maximum entropy approach to modelling species' distributions (Maxent), two generic algorithms based on environmental metrics dissimilarity (BIOCLIM and DOMAIN), Genetic Algorithm for Rule‐set Prediction (GARP), and supervised learning methods based on generalized linear classifiers (support vector machines, SVMs). To test the predictive power of the models we used the Kappa index. Results: Maxent most accurately predicted the habitat distribution of B. balearica, followed by MARS models. The other models tested yielded lower accuracy values. A comparison of the predictive power of the models revealed that climate variables made the highest contributions among the environmental variables studied. The variables that made the lowest contributions were the insolation models. To examine the sensitivity of the models to a reduction in the number of variables, a test showed that accuracy of over 0.90 was maintained by applying just three climatic variables (spring rainfall, mean temperature of the warmest month, and mean temperature of the coldest month). Maps derived from the algorithms of all models tested coincided well with the known distribution of the species. Conclusions: Model habitat prediction is a preliminary step towards highlighting areas of high habitat suitability of B. balearica. These data support the results of previous research, which show that MaxEnt is the best technique for modelling species distributions with small sample sizes.     

Habitat specialization and phylogenetic structure of tree species in a coastal Brazilian white-sand forest

Aims The coastal Brazilian rainforest on white-sand (restinga) ranks among the most fragmented forest types in the tropics, owing to both the patchy distribution of sandy soils and widespread coastal development activities. Here we study the environmental and evolutionary determinants of a forest tree assemblage at a single restinga forest in Southeastern Brazil. We also explore the ability of competing hypotheses to explain the maintenance of species diversity in this forest type, which includes contrasting extremes of edaphic conditions associated with flooding stress.Methods The study was conducted in a white-sand forest permanent plot of 10.24 ha on the coastal plain of Southeastern Brazil. This plot was divided into 256 quadrats of 20×20 m, which were classified into two main edaphic habitats (flooded and drained). Trees with a diameter ≥1cm at breast height were identified. We assembled DNA sequence data for each of the 116 morphospecies recognized using two chloroplast markers (rbcL and matK). A phylogenetic tree was obtained using the maximum likelihood method, and a phylogenetic distance matrix was produced from an ultrametric tree. We analyzed similarity in floristic composition and structure between habitats and related them to cross-plot distances using permutation procedures. Null model torus shift simulations were performed to obtain a statistical significance level for habitat association for each species. The phylogenetic structure for the two habitats and for each 20×20 m quadrat was calculated using the mean phylogenetic distance weighted by species abundance and checked for significance using the standardized effect size generated by 5000 randomizations of phylogenetic tip labels.Important findings Our results indicate that partitioning among edaphic habitats is important for explaining species distributions and coexistence in restinga forests. Species distributions within the plot were found to be non-random: there was greater floristic similarity within than between habitats, and>40% of the more abundant species were positively or negatively associated with at least one habitat. Patterns of habitat association were not independent of phylogenetic relatedness: the community was overdispersed with respect to space and habitat type. Closely related species tended to occur in different habitats, while neighboring trees tended to belong to more distantly related species. We conclude that habitat specialization is important for the coexistence of species in restinga forests and that habitat heterogeneity is therefore an essential factor in explaining the maintenance of diversity of this unique but fragile and threatened type of forest.