Plant diversity,biogeography and environment in Iberia: Patterns and possible causal factors

Abstract. We associated patterns of plant diversity with possible causal factors by considering 93 local regions in the Iberian Peninsula and Balearic Islands with respect to biogeography, environmental favourability, and environmental heterogeneity, and their relationship with measured species diversity at four different scales: mean local species richness standardized at a grain of 100 m², total species richness in a community type within a region (regional community richness), mean compositional similarity, and mosaic diversity. Local regions in biogeographic transition zones to the North African and Atlantic floras had higher regional community richness and greater mosaic diversity than did non‐transitional regions, whereas no differences existed in mean local species richness or mean compositional similarity. Mean local species richness was positively related to environmental favourability as measured by actual evapotranspiration, but negatively related to total precipitation and temporal heterogeneity in precipitation. Mean local species richness was greatest in annual grassland and dwarf shrubland communities, and on calcareous bedrock types. Regional community richness was similarly related to actual evapotranspiration and total precipitation, but in addition was positively related to spatial heterogeneity in topography and soil water holding capacity. Mean compositional similarity decreased with increasing spatial heterogeneity and temperature seasonality. Mosaic diversity, a measure of complexity, increased with increasing local and regional richness. We hypothesize that these relationships can be explained by four ecological and evolutionary classes of causal factors: numbers of individuals, intermediate environments, limits to adaptation, and niche variation. These factors operate at various scales and manifest themselves in various ways. For example, at the site level, apparently processes that increase the number of individuals increase mean local species richness, but at the level of the entire region no such effects were found.     

Patterns of species turnover in plant‐pollinator communities along a precipitation gradient in Patagonia (Argentina)

Recent studies have assessed the influence of different types of gradients (e.g. altitudinal, latitudinal and temporal, among others) on the structure and function of community‐level plant‐pollinator webs. Although the importance of humidity as a major driver of species‐richness gradients worldwide has been stressed by recent reviews, no studies have been specifically designed to address the influence of precipitation gradients on pollination webs. In the present paper we describe for the first time the turnover of species of plants and their associated flower visitors between eight communities located along a steep precipitation gradient in north‐west Patagonia, Argentina. Our results show that: (i) there is a high spatial turnover of plant communities and their associated pollinators; (ii) this turnover is strongly related to precipitation changes across the region; and (iii) the turnover rate is similar for plants and pollinators. Our results support the view that the precipitation gradient is a significant factor associated with the regional turnover of plants and their pollinators in the temperate forests of southern South America.     

Spatial scale, abundance and the species-energy relationship in British birds

1. The spatial scale of analysis may influence the nature, strength and underlying drivers of macroecological patterns, one of the most frequently discussed of which is the relationship between species richness and environmental energy availability. 2. It has been suggested that species-energy relationships are hump-shaped at fine spatial grains and consistently positive at larger regional grains. The exact nature of this scale dependency is, however, the subject of much debate as relatively few studies have investigated species-energy relationships for the same assemblage across a range of spatial grains. Here, we contrast species-energy relationships for the British breeding avifauna at spatial grains of 1 km x 1 km, 2 km x 2 km and 10 km x 10 km plots, while maintaining a constant spatial extent. 3. Analyses were principally conducted using data on observed species richness. While survey work may fail to detect some species, observed species richness and that estimated using nonparametric techniques were strongly positively correlated with each other, and thus exhibit very similar spatial patterns. Moreover, the forms of species-energy relationships using observed and estimated species richness were statistically indistinguishable from each other. 4. Positive decelerating species-energy relationships arise at all three spatial grains. There is little evidence that the explanatory power of these relationships varies with spatial scale. However, ratios of regional (large-scale) to local (small-scale) species richness decrease with increasing energy availability, indicating that local richness responds to energy with a steeper gradient than does regional richness. Local assemblages thus sample a greater proportion of regional richness at higher energy levels, suggesting that spatial turnover of species richness is lower in high-energy regions. Similarly, a crude measure of temporal turnover, the ratio of cumulative species richness over a 4-year period to species richness in a single year, is lower in high-energy regions. These negative relationships between turnover and energy appear to be causal as both total and mean occupancy per species increases with energy. 5. While total density in 1 km x 1 km plots correlates positively with energy availability, such relationships are very weak for mean density per species. This suggests that the observed association between total abundance and species richness may not be mediated by population extinction rates, as predicted by the more individuals hypothesis. 6. The sampling mechanism suggests that species-energy relationships arise as high-energy areas support a greater number of individuals, and that random allocation of these individuals to local areas from a regional assemblage will generate species-energy relationships. While randomized local species-energy relationships are linear and positive, predicted richness is consistently greater than that observed. The mismatch between the observed and randomized species-energy relationships probably arises as a consequence of the aggregated nature of species distributions. The sampling mechanism, together with species spatial aggregation driven by limited habitat availability, may thus explain the species-energy relationship observed at this spatial scale.     

Species richness,vegetation structure,and floristic composition of woody plants along the elevation gradient of Mt. Meru,Tanzania

Understanding the change in vegetation composition along elevational gradients is critical for species conservation in a changing world. We studied the species richness, tree height, and floristic composition of woody plants along an elevation gradient of protected habitats on the eastern slope of Mount Meru and analyzed how these vegetation variables are influenced by the interplay of temperature and precipitation. Vegetation data were collected on 44 plots systematically placed along five transects spanning an elevational gradient of 1600 to 3400 m a.s.l. We used ordinary linear models and multivariate analyses to test the effect of mean annual temperature and precipitation on woody plant species richness, tree height, and floristic composition. We found that species richness, mean tree height, and maximum tree height declined monotonically with elevation. Models that included only mean annual temperature as an explanatory variable were generally best supported to predict changes in species richness and tree height along the elevation gradient. We found significant changes in woody plant floristic composition with elevation, which were shaped by an interaction of mean annual temperature and precipitation. While plant communities consistently changed with temperature along the elevation gradient, levels of precipitation were more important for plant communities at lower than for those at higher elevations. Our study suggests that changes in temperature and precipitation regimes in the course of climate change will reshape elevational gradients of diversity, tree height, and correlated carbon storage in ecosystems, and the sequence of tree communities on East African mountains.     

The relationships between local and regional species richness and spatial turnover

Aim To determine the empirical relationships between species richness and spatial turnover in species composition across spatial scales. These have remained little explored despite the fact that such relationships are fundamental to understanding spatial diversity patterns. Location South‐east Scotland. Methods Defining local species richness simply as the total number of species at a finer resolution than regional species richness and spatial turnover as turnover in species identity between any two or more areas, we determined the empirical relationships between all three, and the influence of spatial scale upon them, using data on breeding bird distributions. We estimated spatial turnover using a measure independent of species richness gradients, a fundamental feature which has been neglected in theoretical studies. Results Local species richness and spatial turnover exhibited a negative relationship, which became stronger as larger neighbourhood sizes were considered in estimating the latter. Spatial turnover and regional species richness did not show any significant relationship, suggesting that spatial species replacement occurs independently of the size of the regional species pool. Local and regional species richness only showed the expected positive relationship when the size of the local scale was relatively large in relation to the regional scale. Conclusions Explanations for the relationships between spatial turnover and local and regional species richness can be found in the spatial patterns of species commonality, gain and loss between areas.     

Dispersal ability links to cross‐scale species diversity patterns across the Eurasian Arctic tundra

Aim The role of dispersal in structuring biodiversity across spatial scales is controversial. If dispersal controls regional and local community assembly, it should also affect the degree of spatial species turnover as well as the extent to which regional communities are represented in local communities. Here we provide the first integrated assessment of relationships between dispersal ability and local‐to‐regional spatial aspects of species diversity across a large geographical area. Location Northern Eurasia. Methods Using a cross‐scale analysis covering local (0.64 m²) to continental (the Eurasian Arctic biome) scales, we compared slope parameters of the dissimilarity‐to‐distance relationship in species composition and the local‐to‐regional relationship in species richness among three plant‐like groups that differ in dispersal ability: lichens with the highest dispersal ability; mosses and moss allies with intermediate dispersal ability; and seed plants with the lowest dispersal ability. Results Diversity patterns generally differed between the three groups according to their dispersal ability, even after controlling for niche‐based processes. Increasing dispersal ability is linked to decreasing spatial species turnover and an increasing ratio of local to regional species richness. All comparisons supported our expectations, except for the slope of the local‐to‐regional relationship in species richness for mosses and moss allies which was not significantly steeper than that of seed plants. Main conclusions The negative link between dispersal ability and spatial species turnover and the corresponding positive link between dispersal ability and the ratio of local‐to‐regional species richness support the idea that dispersal affects community structure and diversity patterns across spatial scales.     

科尔沁沙地固定沙丘植被物种多样性对降水变化的响应

应用１９８６年到１９９２年固定沙丘样方调查数据和同期降水量资料,对科尔沁沙地物种多样性与年降水量的变化进行了分析。结果表明植物种丰富度以对降水量变化的反响比较强烈,一个干旱年份可减少固定沙丘植物种丰富度,而随后的恢复却需要３年友上的时间。物种多样性指数随降水量变化而变化,二者呈正相关。灰色关联分析表明,物种多样性指数Ｄ对降水量变化的反响来看,生长期降水量对物种多样性的影响大于年降水。在固定沙丘植被     

Convergence in growth responses of tropical trees to climate driven by water stress

Widely documented for temperate and cold forests in both hemispheres, variations in tree growth responses to climate along environmental gradients have rarely been investigated in the tropics. Seven tree‐ring chronologies of Centrolobium microchaete (Fabaceae) in the Cerrado tropical forests of Bolivia are used to determine the growth responses to climate along a precipitation gradient. Chronologies are distributed from the humid Guarayos forests (annual precipitation > 1600 mm) in the transition to the Amazonia to the dry‐mesic Chiquitos forests (annual precipitation < 1200 mm) in the proximity to the dry Chaco. On a large spatial scale, radial growth is positively influenced by rainfall and negatively by temperature at the end of the dry season. However, this regional pattern in climate‐tree growth relationship shows differences along the precipitation gradient. Relationships with climate are highly significant and extend over longer periods of the year in sites with low rainfall and extremely severe dry seasons. At wet sites, larger water soil capacity and endogenous forest dynamics partially mask the direct influence of climate on tree growth. Stronger similarities in tree‐growth responses to climate occur between sites in the dry Central Chiquitos and in the transition to the Guarayos forests. In contrast, the relationships show fewer similarities between sites in the humid Guarayos. We conclude that growth responses to climate in the tropics are more similar between sites with limited rainfall and severe and prolonged dry seasons. Our study points to a convergence in the patterns of growth responses of tropical trees to climate, modulated by scarce rainfall and marked seasonality. The negative impact of water deficits on tree physiological processes induces not only the documented reduction in forest species richness, but also a convergence in tree‐growth responses to climate in dry tropical forests.     

Spatiotemporal changes of beetle communities across a tree diversity gradient

Aim Plant and arthropod diversity are often related, but data on the role of mature tree diversity on canopy insect communities are fragmentary. We compare species richness of canopy beetles across a tree diversity gradient ranging from mono‐dominant beech to mixed stands within a deciduous forest, and analyse community composition changes across space and time. Location Germany’s largest exclusively deciduous forest, the Hainich National Park (Thuringia). Methods We used flight interception traps to assess the beetle fauna of various tree species, and applied additive partitioning to examine spatiotemporal patterns of diversity. Results Species richness of beetle communities increased across the tree diversity gradient from 99 to 181 species per forest stand. Intra‐ and interspecific spatial turnover among trees contributed more than temporal turnover among months to the total γ‐beetle diversity of the sampled stands. However, due to parallel increases in the number of habitat generalists and the number of species in each feeding guild (herbivores, predators and fungivores), no proportional changes in community composition could be observed. If only beech trees were analysed across the gradient, patterns were similar but temporal (monthly) species turnover was higher compared to spatial turnover among trees and not related to tree diversity. Main conclusions The changes in species richness and community composition across the gradient can be explained by habitat heterogeneity, which increased with the mix of tree species. We conclude that understanding temporal and spatial species turnover is the key to understanding biodiversity patterns. Mono‐dominant beech stands are insufficient to conserve fully the regional species richness of the remaining semi‐natural deciduous forest habitats in Central Europe, and analysing beech alone would have resulted in the misleading conclusion that temporal (monthly) turnover contributes more to beetle diversity than spatial turnover among different tree species or tree individuals.     

Mean conditions predict salt marsh plant community diversity and stability better than environmental variability

Environmental variability and the frequency of extreme events are predicted to increase in future climate scenarios; however, the role of fluctuations in shaping community composition, diversity and stability is not well understood. Identifying current patterns of association between measures of community stability and climatic means and variability will help elucidate the ways in which altered variability and mean conditions may change communities in the future. Salt marshes provide essential ecosystem services and are increasingly threatened by sea‐level rise, land‐use change, eutrophication and predator loss, yet the effects of temporal environmental variation on salt marshes remain unknown. We synthesized long‐term plant community monitoring data from 11 sites on both coasts of the United States. We used an information‐theoretic approach and linear models to determine the associations among long‐term mean conditions, interannual environmental variability, and plant community stability and diversity. We found that salt marsh community stability and diversity were more strongly related to long‐term means of temperature and precipitation than to interannual variation. Warm and wet environments had fewer species and less turnover among years. Our results suggest that communities in cool, dry environments may be more resilient to climate warming due to greater species richness and turnover. Mean conditions are sufficient to predict contemporary patterns of salt marsh plant community dynamics, but environmental variability may have stronger impacts as it increases with climate change.     

Patterns of floristic richness in vegetation communities of California: regional scale analysis with multi-temporal NDVI

Aim Applying water‐energy dynamics and heterogeneity theory to explain species richness via remote sensing could allow for the regional characterization and monitoring of vegetation community assemblages and their environment. We assess the relationship of multi‐temporal normalized difference vegetation index (NDVI) to plant species richness in vegetation communities. Location California, USA. Methods Sub‐regions containing species inventories for chaparral, coastal sage scrub, foothill woodland, and yellow pine forest communities were intersected with a vegetation community map and an AVHRR NDVI time series for 1990, 1991, 1992, 1995 and 1996. Principal components analysis reduced the AVHRR data to three variables representing the sum and temporal trajectories of NDVI within each community. A fourth variable representing heterogeneity was tested using the standard deviation of the first component. Quadratic forms of these variables were also tested. Species richness was analysed by stepwise regression. Results Chaparral, coastal sage scrub, and yellow pine forest had the best relationships between species richness and NDVI. Richness of chaparral was related to NDVI heterogeneity and spring greenness (r² varied between 0.26 and 0.62 depending on year of NDVI data). Richness of coastal sage scrub was nonlinearly related to annual NDVI and heterogeneity (r² 0.63–0.81), with peak richness at intermediate values. Foothill woodland richness was related to heterogeneity in a monotonic curvilinear fashion (r² 0.28–0.35). Yellow pine forest richness was negatively related to spring greenness and positively related to heterogeneity (r² 0.40–0.46). Main Conclusions While NDVI's relationship to species richness varied, the selection of NDVI variables was generally consistent across years and indicated that spatial variability in NDVI may reflect important patterns in water‐energy use that affect plant species richness. The principal component axis that should correspond closely with annual mean NPP showed a less prominent role. We conclude that plant species richness for coarse vegetation associations can be characterized and monitored at a regional scale and over long periods of time using relatively coarse resolution NDVI data.     

Historical and current climate drive spatial and temporal patterns in fungal endophyte diversity

Horizontally-transmitted foliar endophytic fungi can moderate plant tolerance to abiotic and biotic stress. Previous studies have found correlations between climate and endophyte beta diversity, but were unable to clearly separate drivers related to long-term climate, annual weather, and host plants. To address this, we characterized endophyte communities in the perennial C₄ grass, Panicum hallii, across a precipitation gradient in central Texas over 3 years. A total of 65 unique leaf endophytes were isolated and identified based on ITS and LSU regions of rDNA. Mean annual rainfall and temperature were the primary drivers of endophyte richness and community composition, followed by annual weather conditions. In contrast, little explanatory value was provided by plant host traits, vegetation structure, or spatial factors. The importance of historical climate and annual weather in endophyte distributions suggests that species sort by environment and are likely to be affected by future climate change.     

Liana abundance and diversity increase with rainfall seasonality along a precipitation gradient in Panama

In tropical regions, rainfall gradients often explain the abundance and distribution of plant species. For example, many tree and liana species adapted to seasonal drought are more abundant and diverse in seasonally-dry forests, characterized by long periods of seasonal water deficit. Mean annual precipitation (MAP) is commonly used to explain plant distributions across climate gradients. However, the relationship between MAP and plant distribution is often weak, raising the question of whether other seasonal precipitation patterns better explain plant distributions in seasonally-dry forests. In this study, we examine the relationship between liana abundance and multiple metrics of seasonal and annual rainfall distribution to test the hypothesis that liana density and diversity increase with increasing seasonal drought along a rainfall gradient across the isthmus of Panama. We found that a normalized seasonality index, which combines MAP and the variability of monthly rainfall throughout the year, was a significant predictor of both liana density and species richness, whereas MAP, rainfall seasonality and the mean dry season precipitation (MDP) were far weaker predictors. The strong response of lianas to the normalized seasonality index indicates that, in addition to the total annual amount of rainfall, how rainfall is distributed throughout the year is an important determinant of the hydrological conditions that favor liana proliferation. Our findings imply that changes in annual rainfall and rainfall seasonality will determine the future distribution and abundance of lianas. Models that aim to predict future plant diversity, distribution, and abundance may need to move beyond MAP to a more detailed understanding of rainfall variability at sub-annual timescales.     

Primary Production of Lowland Natural Grasslands and Upland Sown Pastures Across a Narrow Climatic Gradient

Variation in aboveground net primary production (ANPP) is usually studied across wide environmental gradients focusing on spatial averages of zonal natural communities. We studied the spatial and temporal variation of ANPP of upland sown pastures and lowland natural grasslands across a narrow gradient of precipitation and temperature. The Flooding Pampa (Argentina) encompasses an 850–1000 mm range of mean annual precipitation and a 13.8–16.0°C range of mean annual temperature. For 15 100 × 100 km cells, we obtained mean monthly precipitation, temperature, and paddock-level ANPP of upland pastures and lowland grasslands during 8 years. Mean annual ANPP of lowland grasslands and upland sown pastures was positively related to mean annual precipitation. ANPP of upland pastures was 60–80% larger and increased more steeply with mean annual precipitation. ANPP seasonality also changed across the gradient. In lowland grasslands, as mean annual precipitation increased, ANPP monthly maximum increased, minimum decreased, and the duration of the growing season shortened. In contrast, in upland pastures, ANPP monthly maximum was constant, minimum increased, and the growing season lengthened with increasing precipitation. ANPP was more stable across years for lowland grasslands than for upland pastures. The response of annual ANPP to current-year precipitation decreased across the gradient, while the importance of the previous-year precipitation increased. In summary, we found strong spatial and temporal patterns of ANPP across a narrow environmental gradient. In addition, landscape position and species composition heavily influenced those patterns.     

锡林河流域草原群落植物多样性和初级生产力沿水热梯度变化的样带研究

用样带法研究了草原群落植物多样性和初级生产力沿海拔和水分梯度的变化,结果表明;在样带梯度上,物种丰富度、多样性和群落初级生产力与海拔高度、年降水量和土壤有机C及全N含量呈正相关,而与年平均气温和干燥度呈负相关。随着海拔高度的降低,降水量的减少,热量和干燥度的增加,以及土壤有机C和全N含量的降低,草原群落的物种丰富度、多样性和初级生产力逐渐降低。典型相关分析的结果揭示出,土壤有机C含量和干燥度是对草原群落物种丰富度和初级生产力具有更大的影响。同时,草地的利用方式和强度对群落植物多样性和生产力也具有较强的影响。     

Twenty‐five years of plant community dynamics and invasion in New Zealand tussock grasslands

Understanding how plant communities respond to plant invasions is important both for understanding community structure and for predicting future ecosystem change. In a system undergoing intense plant invasion for 25 years, we investigated patterns of community change at a regional scale. Specifically, we sought to quantify how tussock grassland plant community structure had changed and whether changes were related to increases in plant invasion. Frequency data for all vascular plants were recorded on 124, permanent transects in tussock grasslands across the lower eastern South Island of New Zealand measured three times over a period of 25 years. Multivariate analyses of species richness were used to describe spatial and temporal patterns in the vegetation. Linear mixed‐effects models were used to relate temporal changes in community structure to the level and rate of invasion of three dominant invasive species in the genus Hieracium while accounting for relationships with other biotic and abiotic variables. There was a strong compositional gradient from exotic‐ to native‐dominated plant communities that correlated with increasing elevation. Over the 25 years, small‐scale species richness significantly decreased and then increased again; however, these changes differed in different plant communities. Exotic species frequency consistently increased on some transects and consistently declined on others. Species richness changes were correlated with the level of Hieracium invasion and abiotic factors, although the relationship with Hieracium changed from negative to positive over time. Compositional changes were not related to measured predictors. Our results suggest that observed broad‐scale fluctuations in species richness and community composition dynamics were not driven by Hieracium invasion. Given the relatively minor changes in community composition over time, we conclude that there is no evidence for widespread degradation of these grasslands over the last 25 years. However, because of continuing weed invasion, particularly at lower elevations, impacts may emerge in the longer term.     

Plant Functional Traits and the Distribution of West African Rain Forest Trees along the Rainfall Gradient

Plant morphological and physiological traits affect the way plants tolerate environmental stresses and therefore play an important role in shaping species distribution patterns in relation to environmental gradients. Despite our growing knowledge on the role of functional traits in structuring plant communities, few studies have tested their importance at large scales in the wet tropics. Here, we describe the distribution patterns of the most important West African rain forest timber species along the regional rainfall gradient and relate them to their functional traits. We found that the distribution patterns of 25 out of the 31 studied species (80%) were significantly related to mean annual rainfall. Shade tolerance and drought resistance were identified as the main strategy axes of variation. Wood density and leaf deciduousness emerged as the best predictor traits of species position along the rainfall gradient, explaining respectively 32 and 15 percent of the variation. Species traits tended to show stronger relationships with estimated optimum annual rainfall for each species than to the extreme rainfall conditions where they occur. The significant role of rainfall in shaping timber species distribution and the strong relationships between species traits and rainfall indicate that changes in climate, especially declining rainfall, could have strong effects on species composition and abundance in these tropical forests.     

Seasonal Change of Species Diversity Patterns of Non‐volant Small Mammals along Three Subtropical Elevational Gradients

Our understanding of geographic patterns of species diversity and the underlying mechanisms is increasing rapidly, whereas the temporal variation in these patterns remains poorly understood. We examined the seasonal species richness and species turnover patterns of non‐volant small mammals along three subtropical elevational gradients in southwest China. Small mammal diversity was surveyed in two seasons (early wet season and late wet season) using a standardized sampling protocol. The comparison of species richness patterns between two seasons indicated a temporal component in magnitude and shape, with species richness at high elevations clearly increased during the late wet season. Species richness demonstrated weak correlations with modelled temperature and precipitation. The elevational pattern of species turnover measured by Chao‐Sørenson similarity index also changed seasonally, even though the temporal pattern varied with scale. Species turnover between neighboring elevations at high elevations was slower in the late wet season. Meanwhile, there was an acceleration of species turnover along the whole range of the gradient. The seasonal change in species diversity patterns may be due to population‐level increases in abundance and elevational migration, whereas seasonal variation in factors other than temperature and precipitation may play a greater role in driving seasonal diversity patterns. Our study strongly supports the seasonality in elevational patterns of small mammal diversity in subtropical montane forests. Thus it is recommended that subsequent field surveys consider temporal sampling replicate for elevational diversity studies.     

Diversity patterns of stygobiotic crustaceans across multiple spatial scales in Europe

1. Using species distribution data from 111 aquifers distributed in nine European regions, we examined the pairwise relationships between local species richness (LSR), dissimilarity in species composition among localities, and regional species richness (RSR). In addition, we quantified the relative contribution of three nested spatial units – aquifers, catchments and regions – to the overall richness of groundwater crustaceans.
2. The average number of species in karst and porous aquifers (LSR) varied significantly among regions and was dependent upon the richness of the regional species pool (RSR). LSR–RSR relationships differed between habitats: species richness in karstic local communities increased linearly with richness of the surrounding region, whereas that of porous local communities levelled off beyond a certain value of RSR.
3. Dissimilarity in species composition among aquifers of a region increased significantly with increasing regional richness because of stronger habitat specialisation and a decrease in the geographic range of species among karst aquifers. Species turnover among karst aquifers was positively related to RSR, whereas this relationship was not significant for porous aquifers.
4. The contribution of a given spatial unit to total richness increased as size of the spatial unit increased, although 72% of the overall richness was attributed to among-region diversity. Differences in community composition between similar habitats in different regions were typically more pronounced than between nearby communities from different habitats.
5. We conclude by calling for biodiversity assessment methods and conservation strategies that explicitly integrate the importance of turnover in community composition and habitat dissimilarity at multiple spatial scales.