Do vine species in neotropical forests see the forest or the trees?

Questions: Is the occurrence of vine species in neotropical rain forests primarily determined by properties of the forest (environmental factors), by properties of the trees (tree species or tree size) or are vines randomly distributed? Location: Maya Biosphere Reserve, Guatemala. Methods: In five 1‐ha plots that span variation from unlogged forest to forest impacted by recurrent human disturbance we recorded the presence of all climbing vine species on every tree. The presence of all free standing vine species and 11 environmental variables were recorded in 100‐m² subplots. The relationship of host tree diameter and host tree identity on single tree vine species richness was investigated by GLM modelling. Partial redundancy analyses were used to partition the variation in vine species composition on two sources: environmental factors and tree species identity. Results: Single tree vine richness increased with increasing host tree DBH and differed significantly among host species. For climbing vines, the ratio of variation in subplot presence explained by tree species and by environmental variables was ca. 4:1 (in the most disturbed logged plots slightly lower), for free standing vines this ratio varied from 1:2 in the most disturbed logged plots to 9:1 in reserve plots, while a ratio of ca. 1:1 was found for all plots analysed together. Conclusion: Different tree species have different probabilities of being infested by vines. Vines see both the forest and the trees; the environment is more important in earlier developmental stages, properties of individual trees become more important from the time vines start to climb.     

Species richness patterns and metapopulation processes – evidence from epiphyte communities in boreo-nemoral forests

For several epiphyte species, dispersal limitation and metapopulation dynamics have been suggested. We studied the relative importance of local environmental conditions and spatial aggregation of species richness of facultative and obligate epiphytic bryophytes and lichens within two old‐growth forests in eastern Sweden. The effect of the local environment was analyzed using generalized linear models (GLM). We tested whether species richness was spatially structured by fitting variogram models to the residuals of the GLM. In addition, we analyzed the species‐area relationship (area=tree diameter). Different environmental variables explained the richness of different species groups (bryophytes vs lichens, specialists vs generalists, sexual vs asexual dispersal). In most groups, the total variation explained by environmental variables was higher than the variation explained by the spatial model. Spatial aggregation was more pronounced in asexually than in sexually dispersed species. Bryophyte species richness was only poorly predicted by area, and lichen species richness was not explained by area at all. Spatial aggregation may indicate effects of dispersal limitation and metapopulation dynamics on community species richness. Our results suggest that species groups differ in habitat requirements and dispersal abilities; there were indications that presence of species with different dispersal strategies is linked to the age of the host tree. Separate analyses of the species richness of species groups that differ in the degree of habitat specialization and dispersal ability give insights into the processes determining community species richness. The poor species‐area relationship, especially in lichens, may indicate species turnover rather than accumulation during the lifetime of the host tree. Epiphyte species extinctions may be mainly caused by deterministic processes, e.g. changes in habitat conditions as the host tree grows, ages and dies, rather than by stochastic population processes.     

广义模型及分类回归树在物种分布模拟中的应用与比较

比较3个应用较广的模拟物种地理分布模型:广义线性模型(GLM)、广义加法模型(GAM)与分类回归树(CART)对中国树种地理分布模拟的优劣,以提出更为合适的模拟物种地理分布模型,并用于预测气候变化对物种地理分布的影响。3个模型对中国15种树种地理分布的模拟研究表明:除对油松、辽东栎分布的模拟精度稍差外,对其余树种分布的模拟精度均较高,其中以GAM模型最好。结合地理信息系统(GIS),比较分析了这3个模型对青冈、木荷、红松和油松4种树种的地理分布模拟效果,结果亦表明:这3个模型均能很好模拟青冈和木荷的地理分布,而GLM模型对红松分布的模拟结果不太理想,3个模型对油松分布的模拟结果均不甚理想,其中以GLM模型最差。基于3个模型对未来气候变化下青冈与蒙古栎地理分布的预测表明:GLM模型与GAM模型对青冈分布的预测结果较为接近,青冈在未来气候变化情景下向西和向北扩展,而CART模型预测青冈在未来气候变化情景下除有向西、向北扩展趋势外,广东和广西南部的青冈分布区将消失;3个模型均预测蒙古栎在未来气候变化情景下向西扩展,扩展面积的大小为:模型的模拟面积>模型>模型。     

Species–topography association in a species-rich subtropical forest of China

Habitat specialization has been considered as a primary factor in determining the distribution of species. In this study, we investigated species–habitat associations while controlling for spatial neighbourhood effects in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. Habitat specialization was measured by topographic variation and its effects on species distributions were modelled at three different spatial scales (10×10, 20×20 and 25×25 m²) using log-linear regression models and randomization tests. Our results showed: (1) 83% of the species were related to at least one or more topographic variables. Among them, 66%, 60%, 65% and 70% were closely dependent on slope, aspect, elevation and convexity, respectively. (2) Topographic variables have much stronger non-linear (quadratic and cubic) effects on species distributions than linear effects. (3) The effects of topographic heterogeneity on the distribution of shrubs species are smaller than on the distribution of canopy species, and smaller effects were also found in less abundant species. (4) There was a strong neighbourhood effect on species distribution: In 85% of the species, abundance in a focal quadrat was significantly correlated with abundance in the neighbour quadrats. We conclude that habitat specialization plays an important role in maintaining the diversity of this species-rich subtropical forest.     

Environmental Correlates of Tree and Seedling–Sapling Distributions in a Mexican Tropical Dry Forest

Bray and Curtis ordination was used to explore which environmental variables explained importance values and the presence–absence of tropical tree seedlings, saplings and adults in La Escondida-La Cabaña, Sierra de Manantlán, Jalisco, Mexico. The diameters of trees ≥2.5 cm DBH and the presence and height of seedlings and saplings were measured in nine 0.1 ha sites. Four matrices including presence–absence data and importance value indices for trees and seedlings and saplings were analyzed through Bray and Curtis ordination. The matrices were based on density, frequency, and dominance of adult trees as well as seedlings and saplings. The environmental matrix consisted of 18 variables, including elevation, slope, canopy gaps, disturbance, and soil variables. We recorded 63 tree species and 38 seedling and sapling species in the nine sites. The ordination explained 70.9% of the variation in importance value data for trees and 62.6% for seedlings and saplings. The variation explained in presence–absence data for trees was 67.1 and 77.4% for seedlings and saplings. The variance in the ordination axes of seedlings and sapling presence–absence data was poorly explained by the number of gaps in the tree, shrub, or herb layer, suggesting little light specialization by seedlings and saplings. Habitat specialization for soil nutrients appears to be important in explaining the presence–absence of seedlings and saplings. Seedling and sapling specialization along different soil microsites could promote species coexistence in this forest, while heterogeneity in light conditions may instead determine differences in growth and, thus, importance value of trees. We hypothesize that in tropical dry forest in Jalisco, Mexico, a habitat specialization for soil resources is likely more importance at early stages in tree life histories than in later life history.     

Differing Life History Characteristics Support Coexistence of Tree Soil Generalist and Specialist Species in Tropical Rain Forests

Niche differentiation is a prominent mechanism for explaining tree species coexistence in tropical rain forests. Theory predicts that species that are specialized on a set of environmental conditions should competitively exclude generalists in those conditions, and that environmental heterogeneity allows the coexistence of many different species based on niche specialization. Yet, many tropical tree species of the family Dipterocarpaceae have broadly similar habitat preferences, with some occurring widely across several soil types. These soil generalists clearly persist despite the presence of other dipterocarp species that show clear soil specialization. We evaluate comparative seedling performance (growth and survival) of six Shorea species (Dipterocarpaceae) which differ in their adult tree habitat associations within the Sepilok Forest Reserve (Borneo, Malaysia). We tested the hypothesis that seedlings of species associated with a particular soil type perform better on that soil type than seedlings of soil generalist species, with generalists in turn performing better than specialists on a different soil type. We conducted a reciprocal transplant experiment including two soil types (alluvial and mudstone) and two light treatments (gap and understory). The soils differed in soil acidity, Al and P concentration. Observed species‐specific differences in seedling responses to soil, light, and occasional flooding events could partially explain observed adult distribution, although not all species could be fully differentiated along these axes. Other trade‐offs, such as investment in defense against herbivores and tolerance to soil waterlogging, may play additional roles in explaining coexistence of these species.     

Habitat associations of Sterculiaceae trees in a Bornean rain forest plot

Questions: 1. Are trees in a Bornean tropical rain forest associated with a particular habitat? 2. Does the strength of habitat association with the species‐specific optimal habitat increase with tree size? Location : A 52‐ha plot in a mixed dipterocarp forest in a heterogeneous landscape at the Lambir Hills National Park, Sarawak, East Malaysia. Methods : Ten species from the Sterculiaceae were chosen as representative of all species in the plot, on the assumption that competition among closely related species is more stringent than that among more distantly related taxa. Their habitat associations were tested using data from a 52‐ha plot by a torus‐translation test. Results : The torus‐translation test showed that eight out of the ten species examined had significant association with at least one habitat. We could not find negative species‐habitat associations for rare species, probably due to their small sample sizes. Among four species small trees were less strongly associated with habitat than large trees, implying competitive exclusion of trees in suboptimal habitats. The other four species showed the opposite pattern, possibly owing to the smaller sample size of large trees. A habitat had a maximum of three species with which it was significantly positively associated. Conclusions : For a species to survive in population equilibrium in a landscape, habitats in which ‘source’ subpopulations can be sustained without subsidy from adjacent habitats are essential. Competition is most severe among related species whose source subpopulations share the same habitat. On the evidence of source subpopulations identified by positive species‐habitat association, species‐habitat association reduces the number of confamilial competitors. Our results therefore indicate that edaphic niche specialization contributes to coexistence of species of Sterculiaceae in the plot, consistent with the expectations of equilibrium hypotheses.     

Predicting species distributions: a critical comparison of the most common statistical models using artificial species

Aim To test statistical models used to predict species distributions under different shapes of occurrence–environment relationship. We addressed three questions: (1) Is there a statistical technique that has a consistently higher predictive ability than others for all kinds of relationships? (2) How does species prevalence influence the relative performance of models? (3) When an automated stepwise selection procedure is used, does it improve predictive modelling, and are the relevant variables being selected? Location We used environmental data from a real landscape, the state of California, and simulated species distributions within this landscape. Methods Eighteen artificial species were generated, which varied in their occurrence response to the environmental gradients considered (random, linear, Gaussian, threshold or mixed), in the interaction of those factors (no interaction vs. multiplicative), and on their prevalence (50% vs. 5%). The landscape was then randomly sampled with a large (n = 2000) or small (n = 150) sample size, and the predictive ability of each statistical approach was assessed by comparing the true and predicted distributions using five different indexes of performance (area under the receiver‐operator characteristic curve, Kappa, correlation between true and predictive probability of occurrence, sensitivity and specificity). We compared generalized additive models (GAM) with and without flexible degrees of freedom, logistic regressions (general linear models, GLM) with and without variable selection, classification trees, and the genetic algorithm for rule‐set production (GARP). Results Species with threshold and mixed responses, additive environmental effects, and high prevalence generated better predictions than did other species for all statistical models. In general, GAM outperforms all other strategies, although differences with GLM are usually not significant. The two variable‐selection strategies presented here did not discriminate successfully between truly causal factors and correlated environmental variables. Main conclusions Based on our analyses, we recommend the use of GAM or GLM over classification trees or GARP, and the specification of any suspected interaction terms between predictors. An expert‐based variable selection procedure was preferable to the automated procedures used here. Finally, for low‐prevalence species, variability in model performance is both very high and sample‐dependent. This suggests that distribution models for species with low prevalence can be improved through targeted sampling.     

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.     

Long‐term tree fern dynamics linked to disturbance and shade tolerance

Question: Do New Zealand tree ferns have recognizable shade tolerance niches? Location: Lowland temperate rain forest of New Zealand (41°20′S, 174°58′E). Methods: Growth, death and recruitment of five tree fern species were estimated from a 38‐year record of stem heights, collected within a 2.25‐ha block of forest, and electron transport rates (ETR) of photosystem II of fronds were measured. Results: Two species of Cyathea were comparatively common (603 and 351 stems in total) and two were comparatively rare (155 and 17 stems in total) on the site. The common species had lower rates of growth, recruitment and mortality than the rare species, had skewed age distributions typical of shade‐tolerant species and were probably recruited soon after a catastrophic earthquake in 1855. The two rare species were failing to recruit under closed forests; their age distributions indicated that all had regenerated long after the earthquake. ETR were higher for faster‐growing than for the shade‐tolerant species. A tree fern that regenerates vegetatively from aerial buds, Dicksonia squarrosa, was common on the site (361 stems in total). Its age distribution suggested it was relatively shade tolerant, but its mortality and recruitment rates were much higher than those of the two shade‐tolerating Cyathea species, suggesting that this multi‐stemmed species functions differently from the monopodial Cyathea species. Conclusions: New Zealand Cyathea tree ferns occupy distinct niches along a shade tolerance spectrum and their relative abundances are strongly influenced by disturbance history. The study provides evidence that tree fern species differ strongly in their responses to canopy disturbance and are not ecologically equivalent.     

Spatial variation in insect community and species responses to habitat loss and plant community composition

Several experimental studies have examined species responses to manipulations of habitat area and spatial arrangement, but plant composition and spatial variation in species distributions also affect animal responses to habitat alteration. We used an experimental approach to study the combined effects of habitat area, edge, and plant community composition on the spatial structure of insect species richness and composition. The abundance of three guilds (herbivores, predators and parasitoids) and individual species were also analyzed. Habitat patches were created that differed in area and edge by selectively mowing portions of 15 m×15 m plots in a 1.7-ha old field. Spatial and environmental variables were used to predict insect responses in separate multiple regression and ordination models. The variation in species responses due to spatial and environmental variables was then partitioned by combining these variables into an overall regression or ordination. Spatial and environmental variables contributed similar percentages to the total variance in insect species richness, abundance or composition. No significant effects of habitat area were observed in any response variable. Herbivore abundance showed positive responses to legume or grass cover, as well as spatial variation that was unrelated to environmental variables. Predators and parasitoids had greater effects of plant species richness and habitat edge, and less unexplained spatial variation. Individual species differed in their responses to plant variables, depending on host specialization or intraspecific aggregation. Our study highlights the importance of plant community composition and spatial variation apart from environmental variables. Spatial variation stems both from species responses to environmental features as well as species differences in habitat specialization and intraspecific aggregation.     

Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2?years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ??4?years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.     

Vulnerability to xylem embolism as a major correlate of the environmental distribution of rain forest species on a tropical island

Increases in drought‐induced tree mortality are being observed in tropical rain forests worldwide and are also likely to affect the geographical distribution of tropical vegetation. However, the mechanisms underlying the drought vulnerability and environmental distribution of tropical species have been little studied. We measured vulnerability to xylem embolism (P₅₀) of 13 woody species endemic to New Caledonia and with different xylem conduit morphologies. We examined the relation between P₅₀, along with other leaf and xylem functional traits, and a range of habitat variables. Selected species had P₅₀ values ranging between ?4.03 and ?2.00 MPa with most species falling in a narrow range of resistance to embolism above ?2.7 MPa. Embolism vulnerability was significantly correlated with elevation, mean annual temperature and percentage of species occurrences located in rain forest habitats. Xylem conduit type did not explain variation in P₅₀. Commonly used functional traits such as wood density and leaf traits were not related to embolism vulnerability. Xylem embolism vulnerability stands out among other commonly used functional traits as a major driver of species environmental distribution. Drought‐induced xylem embolism vulnerability behaves as a physiological trait closely associated with the habitat occupation of rain forest woody species.     

Heterogeneity in soil water and light environments and dispersal limitation: what facilitates tree species coexistence in a temperate forest?

In the present study, we analysed the habitat association of tree species in an old‐growth temperate forest across all life stages to test theories on the coexistence of tree species in forest communities. An inventory for trees was implemented at a 6‐ha plot in Ogawa Forest Reserve for adults, juveniles, saplings and seedlings. Volumetric soil water content (SMC) and light levels were measured in 10‐m grids. Relationships between the actual number of stems and environmental variables were determined for 35 major tree species, and the spatial correlations within and among species were analysed. The light level had no statistically significant effect on distribution of saplings and seedlings of any species. In contrast, most species had specific optimal values along the SMC gradient. The optimal values were almost identical in earlier life stages, but were more variable in later life stages among species. However, no effective niche partitioning among the species was apparent even at the adult stage. Furthermore, results of spatial analyses suggest that dispersal limitation was not sufficient to mitigate competition between species. This might result from well‐scattered seed distribution via wind and bird dispersal, as well as conspecific density‐dependent mortality of seeds and seedlings. Thus, both niche partitioning and dispersal limitation appeared less important for facilitating coexistence of species within this forest than expected in tropical forests. The tree species assembly in this temperate forest might be controlled through a neutral process at the spatial scale tested in this study.     

Fine‐scale forest variability and biodiversity in the boreal mixedwood forest

Local spatial variation in species distributions is driven by a mix of abiotic and biotic factors, and understanding such hierarchical variation is important for conservation of biodiversity across larger scales. We sought to understand how variation in species composition of understory vascular plants, spiders, and carabid beetles is associated with concomitant spatial variation in forest structure on a 1‐ha permanent plot in a never‐cut mixedwood forest in central Alberta (Canada). Using correlations among dendrograms produced by cluster analysis we associated data about mapped distribution of all living and dead stems > 1 cm diameter at breast height with distributions of the three focal taxa sampled from regular grids across the plot. Variation in each of these species assemblages were significantly associated with several forest structure variables at various spatial scales, but the scale of the associations varied among assemblages. Variation in species richness and abundance was explained mostly by changes in basal area of trees across the plot; however, other variables (e.g. snag density and tree density) were also important, depending on assemblage. We conclude that fine‐scale habitat variation is important in structuring spatial distribution of the species of the forest floor, even within a relatively homogeneous natural forest. Thus, assessments that ignore within‐stand heterogeneity and management that ignores its maintenance will have limited utility as conservation measures for these taxa, which are major elements of forest biodiversity.     

Generalized models vs. classification tree analysis: Predicting spatial distributions of plant species at different scales

Abstract. Statistical models of the realized niche of species are increasingly used, but systematic comparisons of alternative methods are still limited. In particular, only few studies have explored the effect of scale in model outputs. In this paper, we investigate the predictive ability of three statistical methods (generalized linear models, generalized additive models and classification tree analysis) using species distribution data at three scales: fine (Catalonia), intermediate (Portugal) and coarse (Europe). Four Mediterranean tree species were modelled for comparison. Variables selected by models were relatively consistent across scales and the predictive accuracy of models varied only slightly. However, there were slight differences in the performance of methods. Classification tree analysis had a lower accuracy than the generalized methods, especially at finer scales. The performance of generalized linear models also increased with scale. At the fine scale GLM with linear terms showed better accuracy than GLM with quadratic and polynomial terms. This is probably because distributions at finer scales represent a linear sub‐sample of entire realized niches of species. In contrast to GLM, the performance of GAM was constant across scales being more data‐oriented. The predictive accuracy of GAM was always at least equal to other techniques, suggesting that this modelling approach is more robust to variations of scale because it can deal with any response shape.     

Global patterns of specialization and coexistence in bird assemblages

Aim Increased specialization has been hypothesized to facilitate local coexistence and thus high species richness, but empirical evaluations of the richness–specialization relationships have been relatively scant. Here, we provide a first assessment of this relationship for terrestrial bird assemblages at global extent and from fine to coarse grains. Location World‐wide. Methods We use two indices of specialization that describe species‐level resource use: diet and habitat specialization. The relationship between richness and mean assemblage‐level specialization was independently assessed at realm, biome‐realm, 12,100 km² equal‐area grid cells and fine‐grained scales. To identify assemblages that are diverse relative to environmental conditions we: (1) applied quantile regressions, (2) statistically accounted for other environmental variables which may constrain richness, and (3) parsed the data according to the residuals of a model relating species richness to the environmental variables. Results Assemblage species richness increases with both measures of specialization at all scales. Statistically, richness appears constrained by levels of specialization, with the highest richness values only found in specialized assemblages. Richness is positively associated with specialization even after accounting for gradients in resource availability. Net primary productivity and assemblage specialization have complementary statistical effects on assemblage species richness. Contrary to expectations based on niche partitioning of local resources, the relationship between specialization and richness is steep even at coarse scales. Main conclusions The results demonstrate that for an entire clade, totalling > 9000 species, specialization and species richness are related, at least for diverse assemblages. The strong patterns observed across scales suggest that this relationship does not solely originate from (1) limits on coexistence in present‐day assemblages, or (2) increased specialization in richer assemblages imposed by species’ abilities to partition ecological space. Instead, regional‐scale influences on the species pool may determine much of the observed relationship between richness and specialization. Although causal attribution is not straightforward, these findings support the idea that, for the scale of our analysis, specialization may be related to the past origination of high‐diversity assemblages, rather than their contemporary assembly.     

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.     

Species richness–environment relationships within coastal sclerophyll and mesophyll vegetation in Ku‐ring‐gai Chase National Park,New South Wales,Australia

Abstract Patterns in species richness from a wide range of plant communities in Ku‐ring‐gai Chase National Park, New South Wales, Australia, were examined in relation to a number of environmental variables, including soil physical and chemical characteristics. Total species richness and richness of three growth‐form types (trees, shrubs and ground cover) were determined in duplicate 500‐m² quadrats from 50 sites on two geological substrata: Hawkesbury Sandstone and Narrabeen shales and sandstones. Generalized linear models (GLM) were used to determine the amount of variation in species richness that could be significantly explained by the measured environmental variables. Seventy‐three per cent of the variation in total species richness was explained by a combination of soil physical and chemical variables and site attributes. The environmental variables explained 24% of the variation in tree species richness, 67% of the variation in shrub species richness and 62% of the variation in ground cover species richness. These results generally support the hypothesis of an environmental influence on patterns in total species richness and richness of shrubs and ground cover species. However, tree species richness was not adequately predicted by any of the measured environmental variables; the present environment exerts little influence on the richness of this growth‐form type. Historical factors, such as fire or climatic/environmental conditions at time of germination or seedling establishment, may be important in determining patterns in tree species richness at the local scale.     

A Complex Metacommunity Structure for Gastropods Along an Elevational Gradient

The metacommunity framework integrates species‐specific responses to environmental gradients to detect emergent patterns of mesoscale organization. Abiotic characteristics (temperature, precipitation) and associated vegetation types change with elevation in a predictable fashion, providing opportunities to decouple effects of environmental gradients per se from those of biogeographical or historical origin. Moreover, expected structure is different if a metacommunity along an elevational gradient is molded by idiosyncratic responses to abiotic variables (expectation=Gleasonian structure) than if such a metacommunity is molded by strong habitat preferences or specializations (expectation=Clementsian structure). We evaluated metacommunity structure for 13 species of gastropod from 15 sites along an elevational transect in the Luquillo Experimental Forest of Puerto Rico. Analyses were conducted separately for the primary axis and for the secondary axis of correspondence extracted via reciprocal averaging. The metacommunity exhibited quasi‐Clementsian structure along the primary axis, which represented a gradient of gastropod species specialization that was unassociated with elevation. The secondary axis represented environmental variation associated with elevation. Along this axis, the metacommunity exhibited Clementsian structure, with specialists characterizing each of three suites of sites that corresponded to three distinct forest types. These forest types are associated with low (tabonuco forest), mid‐ (palo colorado forest), or high (elfin forest) elevations. Thus, variation among sites in species composition reflected two independent processes: the first decoupled from elevational variation and its environmental correlates, and the second highly associated with environmental variation correlated with elevation. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .