Rarity, commonness, and patterns of species richness: the mammals of Mexico

Aim To determine whether rare or common species contribute most to overall patterns of spatial variation in extant species richness. Location Mexico. Methods Using data on the distribution of mammal species across Mexico at a quarter degree resolution, we ranked species from the most widespread to the most restricted (common‐to‐rare) within the study area, and from the most restricted to the most widespread (rare‐to‐common), and generated a sequence of patterns of species richness for increasing numbers of species. At each stage along both series of richness patterns, we correlated the species richness pattern for the subassemblage with that of the full assemblage. This allows comparison of subassemblages of the n most common with the n most rare species, in terms of how well they match the full assemblage richness pattern. Further analyses examined the effects on these patterns of correlation of the amount of raw information contained in the distributions of given numbers of rare and common species. Results For the mammals of Mexico the more widely distributed species contribute disproportionately to patterns of species richness compared with more restricted species, particularly for non‐volant species and endemic species. This is not simply a consequence of differences in the volumes of information contained in the distributions of rare and common species, with the disproportionate contribution of common species if anything being sharpened when these differences are taken into account. The pattern is most clearly demonstrated by endemic species, suggesting that the contribution of common species is clearest when the causes of rarity and commonness are limited to those genuinely resulting in narrow and widespread geographical ranges, respectively, rather than artificial (e.g. geopolitical) boundaries to the extents of study regions. Conclusions Perhaps surprisingly, an understanding of the determinants of overall patterns of species richness may gain most from consideration of why common species occur in some areas and are absent from others, rather than consideration of the distributions of rare species.     

Reconciling topographic and climatic effects on widespread and range-restricted species richness

Aim To identify the reasons behind differing geographical species richness patterns of range‐restricted and widespread species. Location The Western Hemisphere. Methods We used regression to determine the strongest environmental predictors of richness for widespread and range‐restricted mammal species in 10,000 km² quadrats in the continental Americas. We then used range‐placement models to predict the expected correlation between range‐restricted and widespread species richness were they to be determined by identical, random, or contrasting environmental factors. Finally, to determine the reasons underlying deviations from these predictions, we divided the Americas into 5% quantiles based on temperature and topographic heterogeneity and correlated richness of these two assemblages across quantiles – an approach that avoids constraints on statistical testing imposed by low potential for range overlap among range‐restricted species. Results Minimum annual temperature was the strongest predictor of widespread species richness while topographic heterogeneity was the best, although weak, predictor of range‐restricted species richness in conventional regression analysis. Our models revealed that the observed correlation between range‐restricted and widespread species richness was similar to what would be observed if both range‐restricted and widespread species richness were determined by temperature. Patterns of range‐restricted and widespread species richness were highly correlated across temperature quantiles, but range‐restricted species uniquely showed an increasing pattern across heterogeneity quantiles. Main conclusions Species richness gradients among range‐restricted species differ from those of widespread species, but not as extensively or for the reasons reported previously. Instead, these assemblages appear to share some but not all underlying environmental determinants of species richness. Our new approach to examining species richness patterns reveals that range‐restricted and widespread species richnesses share a common response to temperature that conventional analyses have not previously revealed. However, topographic heterogeneity has assemblage‐specific effects on range‐restricted species.     

The contribution of common and rare species to plant species richness patterns: the effect of habitat type and size of sampling unit

Understanding how overall patterns of spatial variation in species richness are affected by distributional patterns of species has been an area of growing concern. In the present study, we investigated the relative importance of common and rare species as contributors in overall plant species richness. We further examined if the effects of common or rare species in richness patterns are affected by the size of the sampling units and if the observed patterns hold at different habitats. We used a dataset of 5,148 higher plant species distributed across 16,114 sampling plots located in 240 sites of the NATURA 2000 network of Greece. We ranked all species based on the number of sites they occupied and we developed a common to rare and a rare to common sequence. We correlated those sequences with cumulative species distributions. We performed this analysis in nine different sizes of sampling units and in three different datasets referring to (a) all habitat types together, (b) coniferous habitats only and (c) alpine habitats only. Our analysis showed that despite the proportionally higher numbers of restricted species, widespread species make a greater contribution to overall richness patterns and that this observed pattern does not depend on the size of the sampling units. Moreover, the observed pattern stands for different habitat types. Our findings support the generality of this pattern and highlight the importance of widespread species as adequate indicators of biodiversity patterns at various habitat types. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Contribution of rarity and commonness to patterns of species richness

There is little understanding in ecology as to how biodiversity patterns emerge from the distribution patterns of individual species. Here we consider the question of the contributions of rare (restricted range) and common (widespread) species to richness patterns. Considering a species richness pattern, is most of the spatial structure, in terms of where the peaks and troughs of diversity lie, caused by the common species or the rare species (or neither)? Using southern African and British bird richness patterns, we show here that commoner species are most responsible for richness patterns. While rare and common species show markedly different species richness patterns, most spatial patterning in richness is caused by relatively few, more common, species. The level of redundancy we found suggests that a broad understanding of what determines the majority of spatial variation in biodiversity may be had by considering only a minority of species.     

Effects of contemporary environment and Quaternary climate change on drylands plant diversity differ between growth forms

Previous studies on large‐scale patterns in plant richness and underlying mechanisms have mostly focused on forests and mountains, while drylands covering most of the world's grasslands and deserts are more poorly investigated for lack of data. Here, we aim to 1) evaluate the plant richness patterns in Inner Asian drylands; 2) compare the relative importance of contemporary environment, historical climate, vegetation changes, and mid‐domain effect (MDE); and 3) explore whether the dominant drivers of species richness differ across growth forms (woody vs herbaceous) and range sizes (common vs rare). Distribution data and growth forms of 13 248 seed plants were compiled from literature and species range sizes were estimated. Generalized linear models and hierarchical partitioning were used to evaluate the relative contribution of different factors. We found that habitat heterogeneity strongly affected both woody and herbaceous species. Precipitation, climate change since the mid‐Holocene and climate seasonality dominated herbaceous richness patterns, while climate change since the Last Glacial Maximum dominated woody richness patterns. Rare species richness was strongly correlated with precipitation, habitat heterogeneity and historical climatic changes, while common species richness was strongly correlated with MDE (woody) or climate seasonality (herbaceous). Temperature had little effects on the species richness patterns of all groups. This study represents the first evaluation of the large‐scale patterns of plant species richness in the Inner Asian drylands. Our results suggest that increasing water deficit due to anthropogenic activities combined with future global warming may increase the extinction risk of many grassland species. Rare species (both herbaceous and woody) may face severe challenges in the future due to increased habitat destruction caused by urbanization and resource exploitation. Overall, our findings indicate that the hypotheses on species richness patterns based on woody plants alone can be insufficient to explain the richness patterns of herbaceous species.     

Species‐habitat relationships and ecological correlates of butterfly abundance in a transformed tropical landscape

Tropical butterfly conservation strategies often focus on total and/or common species richness to assess the conservation value of a patch or habitat. However, such a strategy overlooks the unique dynamics of rare species. We evaluated the species‐habitat relationships of 209 common, intermediate, and rare butterfly species (including morphospecies) across four habitat types (mature, degraded, or fragmented forest, and urban parks) and two patch sizes (<400 ha, ≥400 ha) in Singapore. Common species richness was consistent across habitat types. Intermediate species richness declined by more than 50 percent in urban parks (relative to all forest habitats), and rare species richness was reduced by 50 percent in degraded and fragmented forest and by 90 percent in urban parks (relative to mature forest). Large patches had comparable overall richness to small patches, but they supported more rare species and three times as many habitat‐restricted species over a similar area. Importantly, a number of rare species were confined to single small patches. Mixed‐effects regression models were constructed to identify habitat and ecological/life history variables associated with butterfly abundance. These models revealed that species with greater habitat specialization, rare larval host plants, few larval host plant genera, and narrow global geographic ranges were more likely to be rare species. Overall, these results demonstrate that the richness of habitat‐restricted and rare species do not follow the same spatial distribution patterns as common species. Therefore, while conserving mature forests is key, effective butterfly conservation in a transformed landscape should take into account rare and habitat‐restricted species.     

Prevalence of intraspecific relationships between range size and abundance in Danish birds

In this study, we investigate patterns in the prevalence of dynamic range–abundance relationships of the Danish avifauna, using breeding bird atlases from 1971 to 1974 and from 1993 to 1996. We focus on differences between common and rare species by dividing the assemblage into range-size quartiles. The trend in total population size was determined using an index. Range was determined as grid cell occupancy and standardized to facilitate comparisons between common and rare species. While narrow-ranging species showed strong and consistent range–abundance relationships, the relationships for widespread species were weak and exhibited considerable variation. This may be due to differences in patterns of resource use, since widespread species generally have wider niches, and so may be less affected by resource-based factors linking range and abundance. Since a tight and dynamic relationship is upheld for rare species, monitoring strategies based on range size surveys seem viable.     

Surface water connectivity drives richness and composition of Arctic lake fish assemblages

相似文献   

Contribution of rare and common species to richness patterns at local scales

Several studies have recently reported that common species are more important for species richness patterns than rare species. However, most such studies have been based on broad‐scale atlas data. We studied the contribution of different species occupancy, i.e. number of plots occupied, to species richness patterns emerging from species data in 50 by 50 m plots within six 140–200 ha forests in Norway. The study included vascular plants, lichens, bryophytes, and polypore fungi. We addressed the following questions: 1) are common species more correlated with species richness than rare species? 2) How do occupancy classes combine at various levels of species richness? 3) Which occupancy class is best in identifying the overall most species‐rich sites (hotspots) by sampling? The results showed that rare species were better correlated with species richness than common species when the information content was accounted for, that high species richness was associated with a higher proportion of less frequent species, and that the best occupancy class for local hotspot identification was species present in 10–30% of the plots within a forest. We argue that the observed correlations between overall richness and sub‐assembly richness are primarily structured by the combination of the distributions of species richness and species occupancy. Although these distributions result from general ecological processes, they may also be strongly affected by idiosyncratic elements of the individual datasets caused by the specific environmental composition of a study area. Hence, different datasets collected in different areas may lead to different results regarding the relative importance of common versus rare species, and such effects should be expected on both broad and fine spatial scales. Despite these effects, we suggest that infrequent species will tend to be more strongly correlated to species richness at local scales than at broader scales as a result of more right‐skewed species‐occupancy distributions.     

Ice age legacies in the geographical distribution of tree species richness in Europe

Aim This study uses a high‐resolution simulation of the Last Glacial Maximum (LGM) climate to assess: (1) whether LGM climate still affects the geographical species richness patterns in the European tree flora and (2) the relative importance of modern and LGM climate as controls of tree species richness in Europe. Location The parts of Europe that were unglaciated during the LGM. Methods Atlas data on the distributions of 55 tree species were linked with data on modern and LGM climate and climatic heterogeneity in a geographical information system with a 60‐km grid. Four measures of species richness were computed: total richness, and richness of the 18 most restricted species, 19 species of medium incidence (intermediate species) and 18 most widespread species. We used ordinary least‐squares regression and spatial autoregressive modelling to test and estimate the richness–climate relationships. Results LGM climate constituted the best single set of explanatory variables for richness of restricted species, while modern climate and climatic heterogeneity was best for total and widespread species richness and richness of intermediate species, respectively. The autoregressive model with all climatic predictors was supported for all richness measures using an information‐theoretic approach, albeit only weakly so for total species richness. Among the strongest relationships were increases in total and intermediate richness with climatic heterogeneity and in restricted richness with LGM growing‐degree‐days. Partial regression showed that climatic heterogeneity accounted for the largest unique variation fraction for intermediate richness, while LGM climate was particularly important for restricted richness. Main conclusions LGM climate appears to still affect geographical patterns of tree species richness in Europe, albeit the relative importance of modern and LGM climate depends on range size. Notably, LGM climate is a strong richness control for species with a restricted range, which appear to still be associated with their glacial refugia.     

Patterns and correlates of plant diversity differ between common and rare species in a neotropical dry forest

Determining which factors affect species richness is important for conservation theory and practice. However, richness of common and rare species may be affected by different factors. We use an extensive inventory of woody plants from a tropical dry forest landscape in Yucatan, Mexico to assess the unique effects of environmental variables, spatial dependence of sampling sites, forest stand age and the combined effect of all groups of variables on species richness of woody plants with different levels of rarity (common, intermediate, rare, very rare)—according to their abundance, habitat specificity and spatial distribution range in the landscape. Analyzing separately common species and those with different levels of rarity uncovered contrasting patterns and correlates of species richness that were not apparent when focusing on all woody plants. In particular, richness of common and intermediate species was influenced mainly by environmental factors, whereas richness of very rare species was affected mostly by the unique effect of spatial dependence of sampling sites, suggesting a main role of environmental filtering and dispersal limitation, respectively. However, common and very rare species also responded inversely to some landscape metrics, revealing contrasting environmental preferences of these groups of species. These contrasting results suggest different underlying mechanisms and the need for very different conservation strategies. Therefore, basic and applied research on tropical forest biodiversity should consider separately species with different levels of rarity, focusing on which factors control variation in each level, and paying special attention to very rare species, generally the most specious and vulnerable to local extinction.     

Native forest metacommunity structures in Uruguay shaped by novel land‐use types in their surroundings

We explore the effect of land‐use change from extensively used grasslands to intensified silvi‐ and agricultural monocultures on metacommunity structure of native forests in Uruguay. We integrated methods from metacommunity studies, remote sensing, and landscape ecology to explore how woody species distribution was influenced by land‐use change from local to regional scale. We recorded richness and composition of adult and juvenile woody species from 32 native forests, created land‐use maps from satellite image to calculate spatial metrics at landscape, class, and patch levels. We also analyzed the influence of land use pattern, climate, topography, and geographic distance between sites (d) on metacommunity, and created maps to visualize species richness and (dis)similarity between communities across the country. Woody species communities were distributed in a discrete pattern across Uruguay. Precipitation and temperature seasonality shaped species distribution pattern. Species richness and community dissimilarity increased from West to East. Latitude did not influence these patterns. Number of patches, landscape complexity, and interspersion and juxtaposition indexes determine woody species distribution at landscape level. Increasing areas covered by crops and timber plantation reduced species richness and increased community dissimilarity. The spatial metrics of native forest fragments at patch level did not influence metacommunity structure, species richness, and community dissimilarity. In conclusion, Uruguayan native forests display a high range of dissimilarity. Pressure of neighborhood land uses was the predominant factor for species assemblages. Conserving landscape structures that assure connectivity within and among native forest patches is crucial. On sites with rare target species, the creation of alliances between governmental institution and landowner complemented by incentives for biodiversity conservation provides opportunities to advance in species protection focused on those less tolerant to land‐use change.     

稀有种和常见种对植物群落物种丰富度格局的相对贡献

物种丰富度格局的形成不仅依赖于群落的构建过程, 同样也依赖于群落中的物种组成(如稀有种和常见种)。本文以黄土高原子午岭林区的辽东栎(Quercus wutaishanica)林为研究对象, 根据频度大小对物种进行排序, 形成稀有-常见种和常见-稀有种两条物种序列, 通过逐一添加(去除)物种, 分析引起的总体物种丰富度及其成分(α多样性和β多样性)的变化, 确定稀有种和常见种对物种丰富度格局的相对贡献。结果表明: (1)常见-稀有种序列与群落总体物种丰富度的相关性呈先剧增后平稳的对数增长曲线, 而稀有-常见种序列与群落总体的相关性与前者刚好相反, 呈先平稳后剧增的指数增长曲线; (2) α多样性在常见-稀有种序列中呈明显的对数变化曲线, 而在稀有-常见种序列中呈指数增长曲线; (3)与α多样性变化相反, β多样性在常见-稀有种序列中随物种的进入先迅速降低后逐渐平稳, 而在稀有-常见种序列中先平稳后急剧降低。可以看出, 常见种不仅主导群落的总体物种丰富度格局, 同时也是α多样性和β多样性格局的重要贡献者。因此, 常见种是群落物种丰富度格局的指示者, 也应该是优先保护的物种。     

Accounting for the capacity of common and rare species to contribute to diversity spatial patterns: Is it a sampling issue or a biological effect?

The contribution of common species to overall species richness in many cases is greater than that of rare species. However, the explanation of this phenomenon remains vague. One hypothesis is that this is a sampling issue and not a biological one. Therefore standardization methods like the information index and empirical variance have been proposed. But, these standardizations do not explicitly compare the significance of the dataset size of the common and rare sub-assemblage. Here, we investigate the role of dataset size in accounting for the capacity of common and rare species to contribute to diversity spatial patterns. We used a dataset of 5148 vascular plant species recorded in 16,439 sample plots in the Greek Natura 2000 network. Species were ranked according to the number of sample plots they occupied in ascending (rare to common), descending (common to rare) and random order. We analyzed the correlation between the richness of each sub-assemblage and total species richness. When comparing among sub-assemblages with equal number of species, common species are clearly the better predictors of total species richness. But, when comparing among sub-assemblages with equal number of occurrence records, the patterns changed. Common and rare species contribution to the overall richness pattern was comparable, with rare species contributing slightly less than widespread species in some cases and the opposite in other cases. However in all cases, sub-assemblages of random species remarkably outperformed the equal sized sub-assemblages of common or rare species. Our results suggest that common and rare species are biased samples of the community and that equal sized random samples are more representative; thus the greater contribution of common species than rare species to biodiversity patterns might be more a sampling issue than a biological effect of commonness or rarity.     

Patterns of rare woody species richness: the influence of environment, landscape attributes and spatial structure across different spatial scales

The aim of this study was to evaluate the relative contributions of the environment, landscape patterns, and spatial structure to explaining the variation in richness of rare woody species at three levels of rarity (low, medium, and high) and at different grain sizes and spatial extents. We used herbarium records of 195 rare woody species to quantify species richness—overall and for three levels of rarity—of the Yucatan Peninsula, Mexico. We assessed relationships between rare species richness and different sets of explanatory variables (environmental, landscape patterns, and spatial structure of sampling units) using linear regression and variation partitioning analyses at three grain sizes (625, 400, and 225 km²). We also conducted a principle coordinates of neighbor matrices analysis to allow interpretation of the results in terms of different spatial extents. The percentage of variation in rare species richness explained by the models was highest for the largest grain size and spatial extent. At the larger extents, rare species richness was explained mainly by the environment, whereas landscape patterns played a more prominent role at the local extent. Landscape patterns also contributed more to explaining species richness at low to medium levels of rarity, whereas the richness of extremely rare species was better explained by spatial structure. We conclude that the relative contribution of the factors explaining the variation of rare species richness depends on both grain and extent, as well as on the level of rarity. These results underscore the importance of considering the different components of scale (grain and extent) as well as different levels of species rarity in order to better understand the patterns of distribution of rare species richness and to be able to frame appropriate conservation strategies.     

Responses of woody vegetation to exclusion of large herbivores in semi‐arid savannas

The Nkuhlu large‐scale long‐term exclusion experiment in Kruger National Park was designed to study the long‐term effects of large herbivores on vegetation. One treatment excludes elephants, another excludes all herbivores larger than hares and another one comprises an open, control area. Vegetation monitoring was implemented in 2002 when a baseline survey was conducted prior to exclusion. Monitoring was repeated 5 years after exclusion. Data from the surveys were analysed to establish how structure and composition of woody vegetation had changed 5 years after herbivore exclusion. The analysis showed that neither plant assemblage nor mean vegetation height had changed significantly since exclusion. However, both species richness and density of woody plants increased 5 years after exclusion of all large herbivores, but not after the exclusion of elephants alone. One already common species, Dichrostachys cinerea, became more common after excluding all large herbivores compared with either no exclusion or elephant exclusion, possibly leading to competitive suppression of other species. Species other than D. cinerea tended to either increase or decrease in density, but the changes were insufficient to induce significant shifts in the overall assemblage of woody plants. The results indicate that after 5 years of exclusion, the combined assemblage of large herbivores, and not elephants alone, could induce changes in species richness and abundances of woody plants, but the effect was so far insufficient to induce measureable shifts in the assemblages of woody plants. It is possible that assemblages will change with time and increasing elephant numbers may amplify future changes.     

Species richness and phylogenetic diversity of seed plants across vegetation zones of Mount Kenya,East Africa

Mount Kenya is of ecological importance in tropical east Africa due to the dramatic gradient in vegetation types that can be observed from low to high elevation zones. However, species richness and phylogenetic diversity of this mountain have not been well studied. Here, we surveyed distribution patterns for a total of 1,335 seed plants of this mountain and calculated species richness and phylogenetic diversity across seven vegetation zones. We also measured phylogenetic structure using the net relatedness index (NRI) and the nearest species index (NTI). Our results show that lower montane wet forest has the highest level of species richness, density, and phylogenetic diversity of woody plants, while lower montane dry forest has the highest level of species richness, density, and phylogenetic diversity in herbaceous plants. In total plants, NRI and NTI of four forest zones were smaller than three alpine zones. In woody plants, lower montane wet forest and upper montane forest have overdispersed phylogenetic structures. In herbaceous plants, NRI of Afro‐alpine zone and nival zone are smaller than those of bamboo zone, upper montane forest, and heath zone. We suggest that compared to open dry forest, humid forest has fewer herbaceous plants because of the closed canopy of woody plants. Woody plants may have climate‐dominated niches, whereas herbaceous plants may have edaphic and microhabitat‐dominated niches. We also proposed lower and upper montane forests with high species richness or overdispersed phylogenetic structures as the priority areas in conservation of Mount Kenya and other high mountains in the Eastern Afro‐montane biodiversity hotspot regions.     

Why does the unimodal species richness–productivity relationship not apply to woody species: a lack of clonality or a legacy of tropical evolutionary history?

Aim  To study how differences in species richness patterns of woody and herbaceous plants may be influenced by ecological and evolutionary factors. Unimodal species richness–productivity relationships (SRPRs) have been of interest to ecologists since they were first described three decades ago for British herbaceous vegetation by J. P. Grime. The decrease in richness at high productivity may be due to competitive exclusion of subordinate species, or diverse factors related to evolution and dispersal. Unimodal SRPRs are most often reported for plants, but there are exceptions. For example, unimodal SRPRs are common in the temperate zone but not in the tropics. Similarly, woody species and forest communities in the Northern Hemisphere do not tend to show unimodal SRPRs.
Location  Global.
Methods  We used data from the literature to test whether a unimodal SRPR applies to woody species and forest communities on a global scale. We explored whether the shape of SRPRs may be related to the lack of clonality in woody species (which may prevent their being competitively superior), or the legacy of evolutionary history (most temperate woody species originate from tropical lineages, and due to niche conservatism they may still demonstrate 'tropical patterns'). We used case studies that reported the names of the dominant or most abundant species for productive sites.
Results  Woody species were indeed less clonal than herbaceous species. Both clonality and the temperate evolutionary background of dominating species were associated with unimodality in SRPRs, with woodiness modifying the clonality effect.
Main conclusions  The unimodal SRPR has been common in the ecological literature because most such studies originate from temperate herbaceous communities with many clonal species. Consequently, both evolutionary and ecological factors may influence species richness patterns.     

Are richness patterns of common and rare species equally well explained by environmental variables?

We investigated relationships between richness patterns of rare and common grassland species and environmental factors, focussing on comparing the degree to which the richness patterns of rare and common species are determined by simple environmental variables. Using data collected in the Machair grassland of the Outer Hebrides of Scotland, we fitted spatial regression models using a suite of grazing, soil physicochemical and microtopographic covariates, to nested sub‐assemblages of vascular and non‐vascular species ranked according to rarity. As expected, we found that common species drive richness patterns, but rare vascular species had significantly stronger affinity for high richness areas. After correcting for the prevalence of individual species distributions, we found differences between common and rare species in 1) the amount of variation explained: richness patterns of common species were better summarised by simple environmental variables, 2) the associations of environmental variables with richness showed systematic trends between common and rare species with coefficient sign reversal for several factors, and 3) richness associations with rare environments: richness patterns of rare vascular species significantly matched rare environments but those of non‐vascular species did not. Richness patterns of rare species, at least in this system, may be intrinsically less predictable than those of common species.     

Relative influence of regional species richness vs local climate on local species richness in China's forests

Disentangling the relative effects of local and regional processes on local species richness (LSR) is critical for understanding the mechanisms underlying large‐scale biodiversity patterns. In this study we used 1098 forest plots from 41 mountains across China, together with regional flora data, to examine the relative influence of local climate vs regional species richness (RSR) on LSR patterns. Both RSR and LSR for woody species and all species combined decreased with increasing latitude, while richness of herbaceous species exhibited a hump‐shaped pattern. The major climatic orrelates of species richness differed across spatial scales. At the regional scale, winter coldness was the best predictor of RSR patterns for both woody and herbaceous species. At the local scale, however, productivity‐related climatic indices were the best predictors of LSR patterns. Local climate and RSR together explained 48, 54 and 23% of the variation in LSR, for overall, woody and herbaceous species, respectively. Both local climate and RSR independently influenced LSR in addition to their joint effects, suggesting that LSR patterns were shaped by local and regional processes together. Local climate and RSR affected LSR of woody species mainly through their joint effects, while there were few shared effects of climate and RSR on the LSR of herbaceous species. Our findings suggest that while geographic RSR patterns are mainly determined by winter coldness, the ecological processes driven by productivity may be critical to the filtering of regional flora into local communities. We also demonstrate that biogeographic region is not a good surrogate for regional richness, at least for our dataset. Consequently, whether biogeographic region can effectively reflect regional effects needs further examination.