Below- and aboveground traits explain local abundance,and regional,continental and global occurrence frequencies of grassland plants

Plants vary widely in how common or rare they are, but whether commonness of species is associated with functional traits is still debated. This might partly be because commonness can be measured at different spatial scales, and because most studies focus solely on aboveground functional traits. We measured five root traits and seed mass on 241 central European grassland species, and extracted their specific leaf area, height, mycorrhizal status and bud-bank size from databases. Then we tested if trait values are associated with commonness at seven spatial scales, ranging from abundance in 16-m² grassland plots, via regional and European-wide occurrence frequencies, to worldwide naturalization success. At every spatial scale, commonness was associated with at least three traits. The traits explained the greatest proportions of variance for abundance in grassland plots (42%) and naturalization success (41%) and the least for occurrence frequencies in Europe and the Mediterranean (2%). Low root tissue density characterized common species at every scale, whereas other traits showed directional changes depending on the scale. We also found that many of the effects had significant non-linear effects, in most cases with the highest commonness-metric value at intermediate trait values. Across scales, belowground traits explained overall more variance in species commonness (19.4%) than aboveground traits (12.6%). The changes we found in the relationships between traits and commonness, when going from one spatial scale to another, could at least partly explain the maintenance of trait variation in nature. Most importantly, our study shows that within grasslands, belowground traits are at least as important as aboveground traits for species commonness. Therefore, belowground traits should be more frequently considered in studies on plant functional ecology.     

Environmental,spatial and phylogenetic determinants of fish life‐history traits and functional composition of Australian rivers

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Functional rarity of plants in German hay meadows — Patterns on the species level and mismatches with community species richness

Functional rarity (FR) — a feature combining a species'' rarity with the distinctiveness of its traits — is a promising tool to better understand the ecological importance of rare species and consequently to protect functional diversity more efficiently. However, we lack a systematic understanding of FR on both the species level (which species are functionally rare and why) and the community level (how is FR associated with biodiversity and environmental conditions). Here, we quantify FR for 218 plant species from German hay meadows on a local, regional, and national scale by combining data from 6500 vegetation relevés and 15 ecologically relevant traits. We investigate the association between rarity and trait distinctiveness on different spatial scales via correlation measures and show which traits lead to low or high trait distinctiveness via distance‐based redundancy analysis. We test how species richness and FR are correlated, and use boosted regression trees to determine environmental conditions that are driving species richness and FR. On the local scale, only rare species showed high trait distinctiveness while on larger spatial scales rare and common species showed high trait distinctiveness. As infrequent trait attributes (e.g., legumes, low clonality) led to higher trait distinctiveness, we argue that functionally rare species are either specialists or transients. While specialists occupy a particular niche in hay meadows leading to lower rarity on larger spatial scales, transients display distinct but maladaptive traits resulting in high rarity across all spatial scales. More functionally rare species than expected by chance occurred in species‐poor communities indicating that they prefer environmental conditions differing from characteristic conditions of species‐rich hay meadows. Finally, we argue that functionally rare species are not necessarily relevant for nature conservation because many were transients from surrounding habitats. However, FR can facilitate our understanding of why species are rare in a habitat and under which conditions these species occur.     

The taxonomic distribution of rare and common species among families in the vascular plant flora of Fennoscandia

Many plant traits are not randomly distributed among families. The question considered here is ‘are rarity and commonness of vascular plants in Fennoscandia randomly distributed among families?’ If more rare or more common species are found within a family, this may give some initial indications about which traits may predict rarity and commonness of species. A species was defined as rare or common based on its abundance and on the number of grid squares it occupies. 1521 naturally occurring species in 229 75×75 km grid squares were used. Permutation tests were performed to assess statistically if rarity and commonness are randomly distributed among families. Several families can be identified as having more rare or more common species than would be expected under a random allocation model. However, there are little deviations from what would be expected if rarity and commonness were randomly distributed among families in the whole Fennoscandian flora. It is proposed that the arbitrary geographical limits of the study area may account for the lack of any clear patterns of rarity and commonness among and between families.     

Growth and reproduction of New Zealand

In New Zealand, as elsewhere, research on rare species has been dominated by autecological studies of individual threatened species. Limitations of this approach are that it involves no comparison with related common species which may have similar traits, and that the minimal sample size prevents generalisation about causes and consequences of rarity. We report on experimentally determined growth and reproductive traits of 10 rare and common Acaena(Rosaceae) species from two taxonomic sections (sect. Ancistrum and sect. Microphyllae). We examined the relationship between rarity or commonness and relative growth rate, mode of vegetative expansion, morphology/presentation of reproductive structures and reproductive allocation. Rarity and commonness were defined according to geographic range size, measured as the number of 10-km grid squares containing at least one record of the species. There were tendencies across both taxonomic sections for species with large range size to have higher relative growth rates and in section Microphyllae, faster lateral expansion. Among section Ancistrum species, common species tended to produce inflorescences for a shorter period and held their capitula higher above the canopy, but other reproductive attributes showed little association with range size. In section Microphyllae all reproductive traits tended to be positively associated with range size. This was mainly due to the single very common species having high fecundity. The lack of strong patterns among our results may reflect insufficient sample size or that the rare species represent different types of rarity.     

Australian Tropical and Subtropical Rain Forest Community Assembly: Phylogeny,Functional Biogeography,and Environmental Gradients

To compare community assemblage patterns in tropical northeastern and subtropical central eastern Australia across selected gradients and scales, we tested the relationship of species traits with phylogenetic structure, and niche breadth. We considered phylogenetic relationships across current‐day species in assemblages in relation to rain forest species pool sizes, and trait values along gradients including elevation and latitude. Trait values were quantified across scales for seed size, leaf area, wood density and maximum height at maturity for 1137 species and 596 assemblages using trait gradient analysis (TGA). Local assemblages of subtropical species had narrower trait ranges, and higher niche breadth values than corresponding assemblages of tropical species. Leaf size and seed size increased at low latitudes, and community phylogenetic structure was most strongly correlated with seed traits in the subtropics, reflecting dispersal and re‐colonization processes. Elevation accounted for little of the variance in community phylogenetic structure or trait variation across local and regional scales. Stable moist forest areas retained many species from ancestral rain forest lineages across a range of temporally conserved habitats; species within assemblages were less related; and rain forest assemblages had higher functional diversity, but lower niche breadth. This suggests that on average, assemblages of species in stable areas had greater trait variation and narrower distributions. Historic and recent rain forest contraction and re‐expansion can result in recolonized areas that are dominated by species that are more related (phylogenetically) than by chance, have smaller, widely dispersed seeds, and greater niche breadth (broader distributions).     

A macroecological perspective of trait patterns in stream communities

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cati: an R package using functional traits to detect and quantify multi‐level community assembly processes

Community ecologists are active in describing species by their functional traits, quantifying the functional structure of plant and animal assemblages and inferring community assembly processes with null‐model analyses of trait distribution and functional diversity indices. Intraspecific variation in traits and effects of spatial scale are potentially important in these analyses. Here, we introduce the R package cati (Community Assembly by Traits: Individuals and beyond) available on CRAN, for the analysis of community assembly with functional traits. cati builds on a recent approach to community assembly that explicitly incorporates individual differences in community assembly analyses and decomposes phenotypic variations across scales and organizational levels, based on three phenotypic variance ratios, termed the T‐statistics. More generally, the cati package 1) calculates a variety of single‐trait and multi‐trait indices from interspecific and intraspecific trait measures; 2) it partitions functional trait variation among spatial and taxonomic levels; 3) it implements a palette of flexible null models for detecting non‐random patterns of functional traits. These patterns can be used to draw inferences about hypotheses of community assembly such as environmental filtering and species interactions. The basic input for cati is a data frame in which columns are traits, rows are species or individuals, and entries are the measured trait values. The cati package can also incorporate a square distance matrix into analyses, which could include phylogenetic or genetic distances among individuals or species. Users select from a variety of functional trait metrics and analyze these relative to a null model that specifies trait distributions in a regional source pool.     

Seven forms of rarity in mammals

Conservation biologists have identified threats to the survival of about a quarter of the mammalian species; to identify patterns of rarity and commonness of mammals, we studied a global sample of 1212 species (about 28% of the mammals) using the ‘7 forms of rarity’ model (in which species are roughly divided into above and below the median for local population density, species’ range area, and number of habitat types). From a niche‐based hypothesis of abundance and distribution, we predicted that mammals would exhibit a bimodal pattern of rarity and commonness, with an overabundance of species in the relatively rarest and most common categories; and just such a significant bimodal pattern emerged, with over a quarter of the species classified as exceedingly rare and a further quarter very common, supporting the niche‐based hypothesis. Orders that include large mammals, including perissodactyls, primates, diprotodonts, and carnivores, exhibited significantly high proportions of relatively rare species; and tropical zoogeographic regions, especially Indomalaya, had relatively high proportions of species in the rarest category. Significant biases in the available data on mammals included under‐sampling of small species like rodents and bats, and a relative paucity of data on zoogeographic regions outside of North America and Australia. Mammalian species listed as of conservation concern by the IUCN occurred in all cells of the model, indicating that even relatively common species can be listed as threatened under some conditions; but we also found that sixty‐three species were relatively rare in all three criteria of the 7‐forms model but were not listed as threatened, indicating potential candidates for further study. Mammals may be a group of animals where rarity or commonness is a natural aspect of species biology, both confirming and perhaps partly explaining the large proportion of mammals assigned threatened status.     

Integrating trait and phylogenetic distances to assess scale‐dependent community assembly processes

Biodiversity is structured by multiple mechanisms that are dependent, at least in part, on ecological similarities and differences among species. Integrating traits and phylogenies in diversity metrics may provide deeper insight into community assembly processes across spatial scales. However, different traits are influenced by processes at different spatial scales, and it is not clear how trait‐spatial scale mismatches skew our ability to detect assembly patterns. An additional complexity is how phylogenetic distances, which might capture unmeasured traits, reflect spatially dependent processes. Here we analyze a freshwater zooplankton dataset from 91 ponds and show that different traits are associated with processes at different spatial scales. We first assessed the response of individual traits to processes at both α‐ and β‐scales, and then quantified the power of different combinations of traits and phylogenetic distances to reveal environmental and spatial drivers of α‐ and β‐diversity. We found that explanatory power was maximised when we accounted for environmental and spatial drivers with single, but different traits for α‐ and β‐diversity. Using the most appropriate trait for each spatial scale outperformed phylogenetic information, but phylogenetic information outperformed the same traits when these were used at the wrong spatial scale, and all outperformed taxonomic analyses that ignore trait and phylogenetic information. We demonstrate that accounting for species’ similarities and differences provides important information about dominant assembly mechanisms at different spatial scales, and that phylogeny is especially useful when measured traits are uninformative at a given spatial scale or when there is lack of trait data. Our study also indicates, however, that trait‐scale mismatches among phylogenetically conserved traits may affect the performance of phylogenetic indices compared to indices that account only for the best single trait at each spatial scale.     

The contribution of geographically common and rare species to the spatial distribution of biodiversity

Motivation and aim

Mapping the spatial distribution of biodiversity is critical for understanding its fundamental drivers (e.g. speciation, environmental filtering) as well as for conservation assessment. An important dimension of this topic is how the distributions of subsets of species contribute to the overall distribution of biodiversity. Although studies have previously investigated the role of geographically common and rare species in determining these patterns, their respective contributions appear to vary between studies. Knowing which species contribute disproportionately to the spatial distribution of biodiversity enables the identification of key indicator species for biodiversity assessments across large areas and is important for prioritising areas for conservation actions. An extensive review of the literature was carried out to synthesise research on how geographic rarity contributes to spatial patterns of biodiversity. We identify potential explanations for the discrepancies in findings between studies and identify opportunities for further research.

Results

Many studies on the contribution of geographic commonness and rarity to the spatial distribution of biodiversity focus on species richness. A prevalent view is that common (widespread) species contribute disproportionately, although this is not ubiquitous across studies due to factors such as the geographic extent from which relative rarity is quantified. We identify research pathways that will further improve our knowledge of how geographically common and rare species shape the spatial distribution of biodiversity including the impact of spatial scale on species contributions and the incorporation of biodiversity components beyond taxonomic alpha diversity, that is functional and phylogenetic diversity.

Main conclusions

Future research should incorporate multiple biodiversity components and model scale dependency. This will further our knowledge on the underlying processes that shape the spatial variation of biodiversity across the planet and help inform biological surveys and conservation activities.     

Scaling up from epidemiology to biogeography: local infection patterns predict geographical distribution in fish parasites

Aim We investigated how the spatial distribution of parasites, measured as either their geographical range size or their frequency of occurrence among localities, relates to either their average local abundance or the variance in their abundance among localities where they occur. Location We used data on the abundance of 46 metazoan parasite species in 66 populations of threespine sticklebacks, Gasterosteus aculeatus, from Europe and North America. Methods For each parasite species, frequency of occurrence was calculated as the proportion of stickleback populations in which it occurred, and geographical range size as the area within the smallest possible polygon delimited using the coordinates of the localities where it occurred. Generalized linear models were used to assess how these two measures of spatial distribution were influenced by several predictor variables: geographical region (North America or Europe), life cycle (simple or complex), average local abundance, the coefficient of variation in abundance across localities, and median prevalence (proportion of infected hosts within a locality). Results Our analyses uncovered four patterns. First, parasites in North America tend to have higher frequencies of occurrence among surveyed localities, but not broader geographical ranges, than those in Europe. Second, parasite species with simple life cycles have wider geographical ranges than those with complex cycles. Third, there was a positive relationship between average abundance of the different parasite species and their frequency of occurrence, but not between average abundance and geographical range size. Fourth, the coefficient of variation in abundance covaried positively with both the frequency of occurrence and geographical range size across the different parasite species. Thus, all else being equal, parasites showing greater site‐to‐site variability in abundance occur in a greater proportion of localities and over a broader geographical area than those with a more stable abundance among sites. Main conclusions Local infection patterns are linked with large‐scale distributional patterns in fish parasites, independently of host effects, such that local commonness translates into regional commonness. The mechanisms linking parasite success at both scales remain unclear, but may include those that maintain the continuum between specialist and generalist parasites. Regardless, the observed patterns have implications for the predicted changes in the geographical distributions of many parasites in response to climate change.     

空间尺度对地形异质性-物种多样性关系的影响:粒度和幅度

空间尺度是影响我们理解生态学格局和过程的关键因素.目前已有多种关于物种多样性分布格局形成机制的假说且研究者未达成共识,原因之一是空间尺度对物种多样性分布格局的环境影响因子的解释力和相对重要性有重要影响.地形异质性是物种多样性分布格局的重要影响因素.本文综述了在地形异质性-物种多样性关系的研究中,不同空间粒度和幅度对研究...     

A phylogenetic and trait‐based analysis of community assembly in a subtropical forest in central China

Despite several decades of study in community ecology, the relative importance of the ecological processes that determine species co‐occurrence across spatial scales remains uncertain. Some of this uncertainty may be reduced by studying the scale dependency of community assembly in the light of environmental variation. Phylogenetic information and functional trait information are often used to provide potentially valuable insights into the drivers of community assembly. Here, we combined phylogenetic and trait‐based tests to gain insights into community processes at four spatial scales in a large stem‐mapped subtropical forest dynamics plot in central China. We found that all of the six leaf economic traits measured in this study had weak, but significant, phylogenetic signal. Nonrandom phylogenetic and trait‐based patterns associated with topographic variables indicate that deterministic processes tend to dominate community assembly in this plot. Specifically, we found that, on average, co‐occurring species were more phylogenetically and functionally similar than expected throughout the plot at most spatial scales and assemblages of less similar than expected species could only be found on finer spatial scales. In sum, our results suggest that the trait‐based effects on community assembly change with spatial scale in a predictable manner and the association of these patterns with topographic variables, indicates the importance of deterministic processes in community assembly relatively to random processes.     

Negative relationship between interspecies spatial association and trait dissimilarity

Species’ response to environmental site conditions and neighborhood interactions are among the important drivers of species’ spatial distributions and the resultant interspecies spatial association. The importance of competition to interspecies spatial association can be inferred from a high degree of trait dissimilarity of the associated species, and vice versa for environmental filtering. However, because the importance of environmental filtering and competition in structuring plant communities often vary with spatial scale and with plant life stage, the species’ spatial association–trait dissimilarity relationship should vary accordingly. We tested these assumptions in a fully mapped 50‐ha subtropical evergreen forest of China, where we assessed the degrees of interspecies spatial associations between adult trees and between saplings at two different spatial scales (10 m versus 40 m) and measured the degrees of trait dissimilarity of the associated species using six traits (leaf area, specific leaf area, leaf dry‐matter content, wood density, wood dry‐matter content and maximum height). Consistent across spatial scales and plant life stages, the degree of interspecies spatial association and the degree of overall trait dissimilarity (i.e. all six traits together) were negatively correlated, suggesting that environmental filtering might help assemble functionally similar species in the forest under study. However, when we looked into the spatial association–trait dissimilarity relationship for individual traits, we found that the relationships between interspecies spatial associations and the dissimilarity of wood density and dry‐matter content were significant for adults but not for saplings, suggesting the importance of wood traits in species’ survival during ontogeny. We conclude that processes shaping interspecies spatial association are spatial scale and plant life stage dependent, and that the distributions of functional traits offer useful insights into the processes underlying community spatial structure.     

Scaling up functional traits for ecosystem services with remote sensing: concepts and methods

Ecosystem service‐based management requires an accurate understanding of how human modification influences ecosystem processes and these relationships are most accurate when based on functional traits. Although trait variation is typically sampled at local scales, remote sensing methods can facilitate scaling up trait variation to regional scales needed for ecosystem service management. We review concepts and methods for scaling up plant and animal functional traits from local to regional spatial scales with the goal of assessing impacts of human modification on ecosystem processes and services. We focus our objectives on considerations and approaches for (1) conducting local plot‐level sampling of trait variation and (2) scaling up trait variation to regional spatial scales using remotely sensed data. We show that sampling methods for scaling up traits need to account for the modification of trait variation due to land cover change and species introductions. Sampling intraspecific variation, stratification by land cover type or landscape context, or inference of traits from published sources may be necessary depending on the traits of interest. Passive and active remote sensing are useful for mapping plant phenological, chemical, and structural traits. Combining these methods can significantly improve their capacity for mapping plant trait variation. These methods can also be used to map landscape and vegetation structure in order to infer animal trait variation. Due to high context dependency, relationships between trait variation and remotely sensed data are not directly transferable across regions. We end our review with a brief synthesis of issues to consider and outlook for the development of these approaches. Research that relates typical functional trait metrics, such as the community‐weighted mean, with remote sensing data and that relates variation in traits that cannot be remotely sensed to other proxies is needed. Our review narrows the gap between functional trait and remote sensing methods for ecosystem service management.     

Effects of soil resource availability on patterns of plant functional traits across spatial scales

Identifying patterns and drivers of plant community assembly has long been a central issue in ecology. Many studies have explored the above questions using a trait‐based approach; however, there are still unknowns around how patterns of plant functional traits vary with environmental gradients. In this study, the responses of individual and multivariate trait dispersions of 134 species to soil resource availability were examined based on correlational analysis and torus‐translation tests across four spatial scales in a subtropical forest, China. Results indicated that different degrees of soil resource availability had different effects on trait dispersions. Specifically, limited resource (available phosphorus) showed negative relationships with trait dispersions, non‐limited resource (available potassium) showed positive relationships with trait dispersions, and saturated resource (available nitrogen) had no effect on trait dispersions. Moreover, compared with the stem (wood density) and architectural trait (maximum height), we found that leaf functional traits can well reflect the response of plants to nutrient gradients. Lastly, the spatial scale only affected the magnitude but not the direction of the correlations between trait dispersions and environmental gradients. Overall, the results highlight the importance of soil resource availability and spatial scale in understanding how plant functional traits respond to environmental gradients.     

Large-scale geographical trends in fruit traits of vertebrate-dispersed temperate plants

Aim  To assess large-scale geographical trends in the character of fleshy, vertebrate-dispersed fruits.
Location  Europe between central Sweden and southern Spain.
Methods  Analyses of fruit of sixty-three plant species from twenty-nine families were compiled from four regional data sets. Four structural and five chemical fruit traits were analysed intraspecifically to control rigorously for phylogenetic lineage effects. Trends were examined in relation to various biological features of the considered species.
Results  Contents of soluble carbohydrate and lipids decreased markedly northwards. Fruit diameter and fresh mass peaked at the wettest site, while the pulp water content remained more constant throughout the gradient than any other fruit trait. Ash content, seed number and seed mass did not change, while the nitrogen content showed conflicting trends. No relation was detected between observed variation in fruit traits and fruit type, fruit colour, ripening season, plant growth form, leaf longevity, or geographical distribution of the considered plant species.
Main conclusions  Considerable intraspecific variability exists in vertebrate-dispersed fruits on large geographical scales. Climate presumably affects particularly those traits related to carbon and water gain and storage. Most research on fruit–frugivore interactions has been carried out on small spatial scales and failed to find matchings between frugivore communities and the character of fleshy fruits. I suggest that explicitly addressed large-scale surveys on the geographical variability of fruits and their disperser assemblages are needed to elucidate their spatial patterns and to determine the extent to which fleshy fruit traits are shaped by animals and/or abiotic factors.     

Geographical expansion and increased prevalence of common species in avian assemblages: implications for large-scale patterns of species richness

Aim The assumption that ecological patterns at large spatial scales originate exclusively from non‐anthropogenic processes is growing more questionable with the increasing domination of the biosphere by humans. Because common and rare species are known to respond differently to anthropogenic activities at local scales these differential responses could, over time, be reflected in distributional patterns of species richness at larger spatial scales. This work tests the hypothesis that modern processes have played a role in shaping these patterns, by examining recent changes in the structure and composition of assemblages of breeding avifauna over a large geographical extent. Location The portion of North America containing the contiguous United States and southern Canada. Methods Changes in the geographical range structure of breeding avifauna in North America from 1968 to 2003 were analysed in regions containing historically moderate levels of anthropogenic activities. Two geographical measures, extent of occurrence and area of occupancy, were used to identify the level of rarity or commonality of individual species and to estimate, based on a vector analysis, patterns of change in geographical range structure for individual species and avian assemblages. Results More species experienced patterns of geographical range expansion (51%) than contraction (28%). The majority of avian assemblages (43%) displayed patterns of geographical range expansion: common species increased in number and proportion (6%) in association with reciprocal losses in rare and moderately rare species, resulting in a constant level of species richness. The minority of avian assemblages (21%) displayed patterns of geographical range contraction: gains occurred for common species as well as for rare and moderately rare species, resulting in substantial increases in species richness and a decline in the proportion of common species (4%). The remaining avian assemblages presented equivocal patterns characterized by gains in the number and proportion (2%) of common species and gains in species richness. Main conclusions Modern processes have played a role in shaping the distribution patterns of species richness at large spatial scales based on the composition of common and rare species. This suggests that anthropogenic activities cannot be ignored as a possible causal factor when considering ecological patterns at large spatial scales.