Tree Diversity,Environmental Heterogeneity,and Productivity in a Mexican Tropical Dry Forest1

Species diversity–environmental heterogeneity (D–EH) and species diversity–productivity (D–P) relationships have seldom been analyzed simultaneously even though such analyses could help to understand the processes underlying contrasts in species diversity among sites. Here we analyzed both relationships at a local scale for a highly diverse tropical dry forest of Mexico. We posed the following questions: (1) are environmental heterogeneity and productivity related?; (2) what are the shapes of D–EH and D–P relationships?; (3) what are individual, and interactive, contributions of these two variables to the observed variance in species diversity?; and (4) are patterns affected by sample size, or by partitioning into average local diversity and spatial species turnover? All trees (diameter at breast height ≥5 cm) within twenty‐six 0.2‐ha transects were censused; four environmental variables associated with water availability were combined into an environmental heterogeneity index; aboveground standing biomass was used as a productivity estimator. Simple and multiple linear and nonlinear regression models were run. Environmental heterogeneity and productivity were not correlated. We found consistently positive log‐linear D–EH and D–P relationships. Productivity explained a larger fraction of among‐transect variance in species diversity than did environmental heterogeneity. No effects of sample size were found. Different components of diversity varied in sensitivity to environmental heterogeneity and productivity. Our results suggest that species' differentiation along water availability gradients and species exclusion at the lowest productivity (driest) sites occur simultaneously, independently, and in a scale‐dependent fashion on the tree community of this forest.     

Habitat and climate heterogeneity maintain beta-diversity of birds among landscapes within ecoregions

Aim  Habitat and climate heterogeneity may affect patterns of species diversity from the relatively local scale of communities to the broad biogeographical scale of continents. However, the effects of heterogeneity on species diversity have not been studied as widely at intermediate scales although differences among landscapes in local climate and habitat should maintain beta-diversity.
Location  Bailey ecoregions in the USA.
Methods  Using a geographically extensive dataset on bird distribution and abundance in 35 ecoregions, we tested for the effects of habitat and climate heterogeneity on beta-diversity at two discrete spatial scales: among sample points within landscapes, and among landscapes within ecoregions.
Results  Landscape-level beta-diversity typically accounted for 50% or more of gamma-diversity and was significantly and positively related to habitat heterogeneity (elevational range within an ecoregion) and climate heterogeneity (variation in potential evapotranspiration). Contrary to predictions, point-level beta-diversity was negatively related to habitat and climate heterogeneity, perhaps because heterogeneity constrains alpha-diversity at the landscape level. The geographical spatial separation of landscapes within an ecoregion did not significantly affect beta-diversity at either scale.
Main conclusions  Our results suggest that habitat selection and adaptation to local climate may be the primary processes structuring bird diversity among landscapes within ecoregions, and that dispersal limitation has a lesser role in influencing beta-diversity among landscapes.     

Scale dependency of processes structuring metacommunities of cladocerans in temporary pools of High‐Andes wetlands

Metacommunity structure can be shaped by a variety of processes operating at different spatial scales. With increasing scale, the compositional variation among local communities (beta diversity) may reflect stronger environmental heterogeneity, but may also reflect reduced exchange of organisms between habitat patches. We analyzed the spatial architecture of a metacommunity of cladoceran zooplankton in temporary pools of High Andes wetlands, with the objective of explaining the spatial dependency of its structure. The spatial distribution of the pools is hierarchical and highly discontinuous: pools are clustered within small wetlands, which lay scattered over valleys that are separated from each other by mountain ridges. We studied a total of 59 pools, belonging to six different wetlands in four different valleys. We assessed pool environmental heterogeneity and sampled active communities and dormant propagule banks of cladoceran zooplankton. Environmental heterogeneity proved very high within wetlands and showed almost no increase with increasing spatial scale. Conversely, diversity partitioning analyses indicated an increase in beta diversity with spatial scale, especially among valleys. Variation partitioning on environmental data and spatial RDA models suggested environmental heterogeneity as the most important generator of beta diversity within wetlands. At the largest spatial scale, beta diversity manifested itself mainly as a differentiation of species occurrence patterns among valleys, which could not be entirely explained by environmental variables. Our study thus presents a case where environmental control seems to be the dominant metacommunity structuring process at the smallest spatial scale, whereas neutral processes and dispersal limitation are the most likely generators of beta diversity at the largest spatial scale.     

Reconceptualising the beta diversity‐environmental heterogeneity relationship in running water systems

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Landscape heterogeneity in relation to variations in epigaeic beetle diversity of a Mediterranean ecosystem. Implications for conservation

Habitat heterogeneity is a determinant cause of biological diversity in natural ecosystems, and therefore its preservation should be a priority when planning conservation strategies. Sierra de Baza, in southern Spain, is a protected natural area in which biotopes of particular interest still remain, but extensive afforestation with pine species has been widespread in recent decades and, in some cases, continues. The aim of this paper is to test the role of habitat heterogeneity at the landscape scale in generating epigaeic beetle diversity in Sierra de Baza. After two-year-long cycles of sampling epigaeic beetles, differences in local diversity at nine sites, and differences in the pattern of species turnover between sites, have been measured. Local communities differed among sites, 74% of the species being scarce (less than 0.41% of total beetle abundance), and thus species replacement differed markedly between sites. Comparisons between habitat types showed that the planted pine forests support less diverse fauna. Our results identify habitat heterogeneity at the landscape scale as the main source of epigaeic beetle diversity at the landscape scale, practices such as extensive afforestation diminishing landscape heterogeneity and consequently local beetle diversity. Preservation of landscape heterogeneity should be encouraged for an adequate beetle biodiversity conservation.     

Heterogeneity fosters biodiversity: Linking history and ecology of dry calcareous grasslands

Calcareous grasslands in Central Europe harbour a high diversity of plant and animal species. However, as man-made habitats, they need to be managed in order to maintain high species diversity. Conservation efforts often aim at reintroducing historical management regimes, such as regular grazing or mowing. Despite such efforts, the diversity and number of species of calcareous grasslands is still decreasing. We propose that, besides fragmentation and eutrophication, a lack of structural heterogeneity within and around calcareous grasslands as created by historic management is causing species loss as well. Here, we review the literature on the history of calcareous grassland management in northern Switzerland, the heterogeneity that it created and the relevance of this heterogeneity for biodiversity at three spatial scales: (1) within grasslands, (2) in their close surroundings and (3) at the landscape scale. Considering that historic management has created heterogeneity at all three scales and that many species do indeed depend on this structural diversity, we conclude that in order to conserve the full range of biodiversity associated with calcareous grasslands, conservation management should aim at increasing heterogeneity in, around, and between grasslands.     

The richness–heterogeneity relationship differs between heterogeneity measures within and among habitats

The positive monotonic relationship between habitat heterogeneity and species richness is a cornerstone of ecology. Recently, it was suggested that this relationship should be unimodal rather than monotonic due to a tradeoff between environmental heterogeneity and population sizes, which increases local species extinctions at high heterogeneity levels. Here, we studied the richness–heterogeneity relationship for an avian community using two different environmental variables, foliage‐height diversity and cover type diversity. We analyzed the richness–heterogeneity within different habitat types (grasslands, savannas, or woodlands) and at the landscape scale. We found strong evidence that both positive and unimodal relationships exist at the landscape scale. Within habitats we found positive relationships between richness and heterogeneity in grasslands and woodlands, and unimodal relationships in savannas. We suggest that the length of the environmental heterogeneity gradient (which is affected by both spatial scale and the environmental variable being analyzed) affects the type of the richness–heterogeneity relationship. We conclude that the type of the relationship between species richness and environmental heterogeneity is non‐ubiquitous, and varies both within and among habitats and environmental variables.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

Beta diversity of stream diatoms at two hierarchical spatial scales: implications for biomonitoring

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Bringing ontology to the gene ontology

We present an analysis of some considerations involved in expressing the Gene Ontology (GO) as a machine-processible ontology, reflecting principles of formal ontology. GO is a controlled vocabulary that is intended to facilitate communication between biologists by standardizing usage of terms in database annotations. Making such controlled vocabularies maximally useful in support of bioinformatics applications requires explicating in machine-processible form the implicit background information that enables human users to interpret the meaning of the vocabulary terms. In the case of GO, this process would involve rendering the meanings of GO into a formal (logical) language with the help of domain experts, and adding additional information required to support the chosen formalization. A controlled vocabulary augmented in these ways is commonly called an ontology. In this paper, we make a modest exploration to determine the ontological requirements for this extended version of GO. Using the terms within the three GO hierarchies (molecular function, biological process and cellular component), we investigate the facility with which GO concepts can be ontologized, using available tools from the philosophical and ontological engineering literature.     

Terminology and quantification of environmental heterogeneity in species‐richness research

Spatial environmental heterogeneity (EH) is an important driver of species diversity, and its influence on species richness has been analysed for numerous taxa, in diverse ecological settings, and over a large range of spatial scales. The variety and ambiguity of concepts and terminology, however, have hampered comparisons among studies. Based on a systematic literature survey of 192 studies including 1148 data points, we provide an overview of terms and measures related to EH, and the mechanisms that relate EH to species richness of plants and animals in terrestrial systems. We identify 165 different measures used to quantify EH, referred to by more than 350 measure names. We classify these measures according to their calculation method and subject area, finding that most studies have analysed heterogeneity in land cover, topography, and vegetation, whereas comparatively few studies have focused on climatic or soil EH. Overall, elevation range emerged as the most frequent measure in our dataset. We find that there is no consensus in the literature about terms (such as ‘habitat diversity’ or ‘habitat complexity’), their meanings and associated quantification methods. More than 100 different terms have been used to denote EH, with largely imprecise delimitations. We reveal trends in use of terms and quantification with respect to spatial scales, study taxa, and locations. Finally, we discuss mechanisms involved in EH–richness relationships, differentiating between effects on species coexistence, persistence, and diversification. This review aims at guiding researchers in their selection of heterogeneity measures. At the same time, it shows the need for precise terminology and avoidance of ambiguous synonyms to enhance understanding and foster among‐study comparisons and synthesis.     

Biological races in humans

Races may exist in humans in a cultural sense, but biological concepts of race are needed to access their reality in a non-species-specific manner and to see if cultural categories correspond to biological categories within humans. Modern biological concepts of race can be implemented objectively with molecular genetic data through hypothesis-testing. Genetic data sets are used to see if biological races exist in humans and in our closest evolutionary relative, the chimpanzee. Using the two most commonly used biological concepts of race, chimpanzees are indeed subdivided into races but humans are not. Adaptive traits, such as skin color, have frequently been used to define races in humans, but such adaptive traits reflect the underlying environmental factor to which they are adaptive and not overall genetic differentiation, and different adaptive traits define discordant groups. There are no objective criteria for choosing one adaptive trait over another to define race. As a consequence, adaptive traits do not define races in humans. Much of the recent scientific literature on human evolution portrays human populations as separate branches on an evolutionary tree. A tree-like structure among humans has been falsified whenever tested, so this practice is scientifically indefensible. It is also socially irresponsible as these pictorial representations of human evolution have more impact on the general public than nuanced phrases in the text of a scientific paper. Humans have much genetic diversity, but the vast majority of this diversity reflects individual uniqueness and not race.     

Measuring biological heterogeneity of forest vegetation types: avalanche index as an estimate of biological diversity

Ideally, the estimates of biological diversity of a community of species in a habitat should refer to the biological variation among the species and not merely to their numbers and frequencies. However, the current estimates of biodiversity incorporate only the latter two components but not the biological differences among the species. Ganeshaiah et al. [(1997) Current Science 73: 128–133] have proposed an estimate called the Avalanche Index (AI) that can incorporate the biological heterogeneity among the species in a habitat. This estimate, besides being methodologically simple, can incorporate any quantifiable differences among the species, information on species richness and their frequencies in the habitat. In this paper we have estimated AI for tree vegetation in 14 forest types across different ecosystems of the world and have compared these estimates with other indices being currently used. Through this we have attempted to analyse the relative utility of AI in discriminating the habitats based on their biological heterogeneity by capturing their intra-community biological variation. We discuss the merits and demerits of the AI as a comprehensive estimate of biological diversity.     

The latitudinal gradient of species diversity among North American grasshoppers (Acrididae) within a single habitat: a test of the spatial heterogeneity hypothesis

Abstract. The spatial heterogeneity hypothesis predicts a positive relationship between habitat complexity and species diversity: the greater the heterogeneity of a habitat, the greater the number of species in that habitat. On a regional scale, this hypothesis has been proposed to explain the increases in species diversity from the poles to the tropics: the tropics are more diverse because they contain more habitats. On the local scale, the spatial heterogeneity hypothesis suggests that the tropics are more diverse because they contain more microhabitats. The positive relationship between habitat heterogeneity and species diversity, on the local scale, is well documented. In this paper, we test whether habitat heterogeneity on the local scale can explain the latitudinal gradient of species diversity on the regional scale. We determined the latitudinal gradient of species diversity of 305 species of North American grasshoppers using published distribution maps. We compared the slope of this multihabitat (regional-scale) gradient with the slope of a within-habitat (local-scale) gradient in the prairie grasslands. Our results show no significant difference between the slopes at the two scales. We tested the generality of our results by comparing multi- and within-habitat latitudinal gradients of species diversity for ants, scorpions and mammals using data from the literature. These results are in accordance with those from grasshoppers. We can therefore reject the local-scale spatial heterogeneity hypothesis as a mechanism explaining the regional-scale latitudinal gradient of species diversity. We discuss alternative mechanisms that produce this gradient.     

Microtopographic heterogeneity constrains alpine plant diversity, Glacier National Park, MT

Theoretical and empirical evidence exists for a positive relationship between environmental heterogeneity and species diversity. Alpine plant communities can exhibit exceptional diversity at a fine scale, which niche theory would suggest is the result of fine scale spatial heterogeneity of the environment. To test if species diversity of alpine plants is driven by environmental heterogeneity, we sampled vascular plant species composition, microtopography, and ground cover within 1?m² plots with and without solifluction forms in Glacier National Park, MT. We analyzed the relationship between microtopographic heterogeneity and species richness at the plot and sub-plot scale with linear and quantile regression, respectively. Species richness does not differ between the plots varying in cover type. Species richness is negatively related to the fractal dimension (D) of the ground surface and non-vegetated ground cover within 1?m² plots. At a finer scale, the standard deviation of elevation and slope appear to impose a limit on species richness such that more variable sub-plots have lower species richness. Contrary to our expectations, microtopographic heterogeneity does not promote the diversity of alpine plants. The negative relationship between topographic heterogeneity and species richness is contrary to the theoretical prediction that environmental heterogeneity generally results in greater species diversity. It is possible that microtopographic variability represents a measure of soil disturbance, which would be expected to have a negative effect on species diversity in alpine tundra due to its low productivity.     

Spatial pattern of distribution of marine invertebrates within a subtidal community: do communities vary more among patches or plots?

Making links between ecological processes and the scales at which they operate is an enduring challenge of community ecology. Our understanding of ecological communities cannot advance if we do not distinguish larger scale processes from smaller ones. Variability at small spatial scales can be important because it carries information about biological interactions, which cannot be explained by environmental heterogeneity alone. Marine fouling communities are shaped by both the supply of larvae and competition for resources among colonizers—these two processes operate on distinctly different scales. Here, we demonstrate how fouling community structure varies with spatial scale in a temperate Australian environment, and we identify the spatial scale that captures the most variability. Community structure was quantified with both univariate (species richness and diversity) and multivariate (similarity in species composition) indices. Variation in community structure was unevenly distributed between the spatial scales that we examined. While variation in community structure within patch was usually greater than among patch, variation among patch was always significant. Opportunistic taxa that rely heavily on rapid colonization of free space spread more evenly among patches during early succession. In contrast, taxa that are strong adult competitors but slow colonizers spread more evenly among patches only during late succession. Our findings show significant patchiness can develop in a habitat showing no systematic environmental spatial variation, and this patchiness can be mediated through different biological factors at different spatial scales.     

Experimental evidence that soil heterogeneity enhances plant diversity during community assembly

Aims Environmental heterogeneity is a primary mechanism explaining species coexistence and extant patterns of diversity. Despite strong theoretical support and ample observational evidence, few experimental studies in plant communities have been able to demonstrate a causal link between environmental heterogeneity and plant diversity. This lack of experimental evidence suggests that either fine-scale heterogeneity has weak effects on plant diversity or previous experiments have been unable to effectively manipulate heterogeneity. Here, we utilize a unique soil manipulation to test whether fine-scale soil heterogeneity will increase plant richness through species sorting among experimental patch types.Methods This experiment was conducted in the tallgrass prairie region of south-central Kansas, USA. We utilized the inherent variation found in the vertical soil profile, which varied in both biotic and abiotic characteristics, and redistributed these strata into either homogeneous or heterogeneous spatial arrangements in 2.4×2.4 m plots. After the soil manipulation, 34 native prairie species were sown into all plots. We conducted annual censuses at peak biomass to quantify species composition and plant density by species within the experimental communities.Important findings After 2 years, species richness was significantly higher in heterogeneous relative to homogeneous plots and this pattern was independent of total plant density. In the heterogeneous plots, 13 species had higher establishment in a specific patch type representing one of the three soil strata. Conversely, no species had greater establishment in the mixed stratum, which comprised the homogeneous plots, relative to the heterogeneous strata. These species sorting patterns suggest that fine-scale heterogeneity creates opportunities for plant establishment due to niche differences, which translates into increased plant diversity at the plot scale. Species richness was more strongly related to plant density among patches comprising homogenous plots—where fine-scale heterogeneity was minimized, but weak in heterogeneous plots. This pattern is consistent with the idea that richness–density relationships dominate when neutral processes are important but are weak when niche processes operate. Unlike many previous attempts, our results provide clear, experimental evidence that fine-scale soil heterogeneity increases species richness through species sorting during community assembly.     

The geometry of coexistence

Understanding the processes that maintain diversity has been the focus of extensive study, yet there is much that has not been integrated into a cohesive framework. First, there is a separation of perspective. Ecological and evolutionary approaches to diversity have progressed in largely parallel directions. Second, there is a separation of emphasis. In both ecology and population genetics, classical theories favour local explanations with emphasis on population dynamics and selection within populations, while contemporary theories favour spatial explanations, with emphasis on population structure and interactions among populations. What is lacking is a comparative approach that evaluates the relative importance of local and spatial processes in maintaining genetic and ecological diversity. I present a framework for diversity maintenance that emphasizes the comparative approach. I use a well-known but little-used mathematical approach, the perturbation theorem for dynamical systems, to identify key points of contact between ecological and population genetic theories of coexistence. These connections provide for a synthesis of several important concepts: population structure (source-sink versus extinction-colonization), spatial heterogeneity (intrinsic versus extrinsic) in fitness and competitive ability, and temporal scales over which local and spatial processes influence diversity. This framework ties together a large and diverse body of theory and data from ecology and population genetics. It yields comparative predictions that can serve as guidelines in biodiversity management.     

Genetic and pharmacological aspects of histamine H3 receptor heterogeneity

Histaminergic H3 receptors modulate the release of neurotransmitters within the CNS and periphery. Ligands for these receptors have potential clinical utility in a variety of disease states. However, the pharmacological characteristics of these receptors have been enigmatic for more than a decade because of the diversity of pharmacological effects observed with the limited number of heretofore-available compounds. Recent cloning of the H3 receptor has revealed interspecies differences in the protein sequences in key regions, the existence of splice variants that differ in composition between species, and potential differences in signal transduction processes between either different tissues and/or species. This review attempts to summarize these findings within the context of the molecular biological and pharmacological data accumulated to date. Also, we suggest a nomenclature strategy to reduce potential confusion that has arisen from different naming systems used by various investigators. While some facets of this genetic and pharmacological diversity help to rationalize various aspects of H3 receptor heterogeneity, there remains an insufficient repertoire of selective ligands, assays, or other measures to completely resolve all components of this diversity. The promise of newly available tools to further explore H3 receptor function may provide the insight to bring the promised clinical potential of H3 receptor ligands to realization.     

The intermediate disturbance hypothesis does not explain fire and diversity pattern in fynbos

The intermediate disturbance hypothesis is a widely accepted generalization regarding patterns of species diversity, but may not hold true where fire is the disturbance. In the Mediterranean-climate shrublands of South Africa, called fynbos, fire is the most importance disturbance and a controlling factor in community dynamics. The intermediate disturbance hypothesis states that diversity will be highest at sites that have had an intermediate frequency of disturbance and will be lower at sites that have experienced very high or very low disturbance frequencies. Measures of diversity are sensitive to scale; therefore, we compared species richness for three fire regimes in South African mountain fynbos to test the intermediate disturbance hypothesis over different spatial scales from 1 m² to 0.1 hectares. Species diversity response to fire frequency was highly scale-dependent, but the relationship between species diversity and disturbance frequency was opposite that predicted by the intermediate disturbance hypothesis. At the largest spatial scales, species diversity was highest at the least frequently burned sites (40 years between fires) and lowest at the sites of moderate (15 to 26 years between fires) and high fire frequency (alternating four and six year fire cycle). Community heterogeneity, measured both as the slope of the species-area curve for a site and as the mean dissimilarity in species composition among subplots within a site, correlated with species diversity at the largest spatial scales. Community heterogeneity was highest at the least frequently burned sites and lowest at the sites that experienced an intermediate fire frequency.