Richness gradients of stream invertebrates across the USA: taxonomy- and trait-based approaches

Large-scale diversity patterns in relationship to environmental factors at multiple spatial scales have been well-studied for many taxonomic groups; however, freshwater ecosystems remain understudied. Biodiversity is now widely recognized to encompass many more factors than just species numbers, particularly the inclusion of functional attributes. In this study, we examined richness patterns of stream invertebrate genera and their biological traits (“functional” richness) across 364 sites in the contiguous USA. In particular, we focused on the relationship between taxonomy- and trait-based richness to test for functional redundancy in stream communities. Further, we obtained environmental data to model the relative importance of local and watershed-scale environmental factors and residual spatial (latitude, longitude) influences on taxonomy- and trait-based richness. Trait richness increased linearly with genus richness (slope ≪ 1), although this appears to be an artifact of the restricted range of genus richness in our study (32 genera maximum). Furthermore, trait richness was significantly lower than expected under random community assembly. In contrast, the Ephemeroptera, Plecoptera, and Trichoptera (EPT) genera exhibited a saturating pattern between trait and genus richness and trait richness was no different from random. Our study indicates that there is functional redundancy among stream invertebrate genera, likely as a result of harsh habitat filters limiting trait diversity. Environmental factors (including spatially structured environmental factors) were always more important than spatial factors (latitude, longitude) in structuring richness despite strong longitudinal patterns of all richness measures (these differences were only significant for EPT genera). Finally, we found no significant difference in the relative importance of local and watershed scale environmental factors for taxonomy- and trait-based richness.     

A functional trait-based approach to understand community assembly and diversity–productivity relationships over 7 years in experimental grasslands

Several multi-year biodiversity experiments have shown positive species richness–productivity relationships which strengthen over time, but the mechanisms which control productivity are not well understood. We used experimental grasslands (Jena Experiment) with mixtures containing different numbers of species (4, 8, 16 and 60) and plant functional groups (1–4; grasses, legumes, small herbs, tall herbs) to explore patterns of variation in functional trait composition as well as climatic variables as predictors for community biomass production across several years (from 2003 to 2009). Over this time span, high community mean trait values shifted from the dominance of trait values associated with fast growth to trait values suggesting a conservation of growth-related resources and successful reproduction. Increasing between-community convergence in means of several productivity-related traits indicated that environmental filtering and exclusion of competitively weaker species played a role during community assembly. A general trend for increasing functional trait diversity within and convergence among communities suggested niche differentiation through limiting similarity in the longer term and that similar mechanisms operated in communities sown with different diversity. Community biomass production was primarily explained by a few key mean traits (tall growth, large seed mass and leaf nitrogen concentration) and to a smaller extent by functional diversity in nitrogen acquisition strategies, functional richness in multiple traits and functional evenness in light-acquisition traits. Increasing species richness, presence of an exceptionally productive legume species (Onobrychis viciifolia) and climatic variables explained an additional proportion of variation in community biomass. In general, community biomass production decreased through time, but communities with higher functional richness in multiple traits had high productivities over several years. Our results suggest that assembly processes within communities with an artificially maintained species composition maximize functional diversity through niche differentiation and exclusion of weaker competitors, thereby maintaining their potential for high productivity.     

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain,Southwest China

Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.     

Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages

Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co‐occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within‐community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities.     

Dominance determines fish community biomass in a temperate seagrass ecosystem

Biodiversity and ecosystem function are often correlated, but there are multiple hypotheses about the mechanisms underlying this relationship. Ecosystem functions such as primary or secondary production may be maximized by species richness, evenness in species abundances, or the presence or dominance of species with certain traits. Here, we combine surveys of natural fish communities (conducted in July and August 2016) with morphological trait data to examine relationships between biodiversity and ecosystem function (quantified as fish community biomass) across 14 subtidal eelgrass meadows in the Northeast Pacific (54°N, 130°W). We employ both taxonomic and functional trait measures of diversity to investigate whether ecosystem function is best predicted by species diversity (complementarity hypothesis) or by the presence or dominance of species with particular trait values (selection or dominance hypotheses). After controlling for environmental variation, we find that fish community biomass is maximized when taxonomic richness and functional evenness are low, and in communities dominated by species with particular trait values, specifically those associated with benthic habitats and prey capture. While previous work on fish communities has found that species richness is often positively correlated with ecosystem function, our results instead highlight the capacity for regionally prevalent and locally dominant species to drive ecosystem function in moderately diverse communities. We discuss these alternate links between community composition and ecosystem function and consider their divergent implications for ecosystem valuation and conservation prioritization.     

The ecology of differences: assessing community assembly with trait and evolutionary distances

Species enter and persist in local communities because of their ecological fit to local conditions, and recently, ecologists have moved from measuring diversity as species richness and evenness, to using measures that reflect species ecological differences. There are two principal approaches for quantifying species ecological differences: functional (trait‐based) and phylogenetic pairwise distances between species. Both approaches have produced new ecological insights, yet at the same time methodological issues and assumptions limit them. Traits and phylogeny may provide different, and perhaps complementary, information about species' differences. To adequately test assembly hypotheses, a framework integrating the information provided by traits and phylogenies is required. We propose an intuitive measure for combining functional and phylogenetic pairwise distances, which provides a useful way to assess how functional and phylogenetic distances contribute to understanding patterns of community assembly. Here, we show that both traits and phylogeny inform community assembly patterns in alpine plant communities across an elevation gradient, because they represent complementary information. Differences in historical selection pressures have produced variation in the strength of the trait‐phylogeny correlation, and as such, integrating traits and phylogeny can enhance the ability to detect assembly patterns across habitats or environmental gradients.     

Disturbance regimes in a wetland remnant: implications for trait-displacements and shifts in the assemblage structure of carabid beetles (Coleoptera: Carabidae)

Studies on disturbance regimes involving carabid beetles have mainly focused on forest habitats. We therefore decided to analyze the effects of disturbance on carabid communities in a wetland remnant (Lake Chiusi, central Italy). Results highlighted the presence of a disturbance gradient affecting the species richness and trait-displacement of carabid communities. Carabids were sampled with pitfall traps from March to October 2008 at nine randomly selected sample stations; a set of landscape attributes were also collected. Principal Component Analysis (PCA) and generalized linear mixed models (GLMMs) were used to link the distribution of carabid life-history traits and species richness with the most informative combination of landscape attributes. The first PCA axis (PC1) showed significant correlation with “distance to the lake shoreline” and “perimeter-area ratio”, highlighting the presence of a disturbance-axis. The second and third axes accounted only for a trivial portion of the total variance. GLMMs revealed a progressive decrease in the number of hygrophilous species from the core of the wetland to its outer areas. Similar trends were observed for species richness and for predator species with good dispersal ability and larval period in summer. Our results highlight the importance of taking into account community-wide functional implications in landscape ecology studies.     

Leaf trait integration mediates species richness variation in a species-rich neotropical forest domain

Environmental gradients are known to drive changes in mean trait values, but changes in the trait integration strength across local communities are less well understood, particularly with regard to possible links with species richness variation. Here, we tested if climate, soil, and topography gradients drive species richness indirectly via constraints on trait integration in the Atlantic Forest of South America. We evaluated seven traits (from leaf, wood, seed, and plant size) of 1456 species occurring across 84 local communities. Generalized least square models and a path model were applied to test direct and indirect relationships. Correlations were higher between leaf traits (average r?=?0.28) and lower when other traits were included (average r?=?0.16). In line with this result, species richness was related to a multivariate index of interspecific trait integration (ITI) computed for leaf traits, but not to the ITI for all the seven traits. Abiotic gradients influenced species richness both directly and indirectly through the leaf trait integration. A total of 33% and 26% of the variation in species richness and ITI, respectively, were explained by the models, with climatic conditions showing higher contribution than topographic and edaphic factors. These results support a significant but reduced environmental selection role behind the trait-based community assembly and may suggest that other processes are involved in the constrain of trait integration at larger spatial scales. In addition, different directional trends in trait–trait relationships across local communities suggest that global trait relationships may not necessarily hold at local contexts.

     

CWM and Rao’s quadratic diversity: a unified framework for functional ecology

Assessing the effects of environmental constraints on community structure often relies on methods that consider changes in species functional traits in response to environmental processes. Various indices have been proposed to measure relevant aspects of community trait composition from different viewpoints and perspectives. Among these, the ‘community-weighted mean trait value’ (CWM) and the Rao coefficient have been widely used in ecological research for summarizing different facets of functional composition and diversity. Analyzing changes in functional diversity of bee communities along a post-fire successional gradient in southern Switzerland we show that these two measures may be used to describe two complementary aspects of community structure, such as the mean and the dispersion of functional traits within a given species assemblage. While CWM can be adequately used to summarize shifts in mean trait values within communities due to environmental selection for certain functional traits, the Rao coefficient can be effectively applied to analyze patterns of trait convergence or divergence compared to a random expectation.     

Functional traits,spatial patterns and species associations: what is their combined role in the assembly of wetland plant communities?

Studies of community assembly focus on finding rules that predict which species can become member of a plant community. Within a community, species can be categorized in two ways: functional groups classify species according to their functional traits, whereas generalized guilds group species based on their (co-)occurrence, spatial distribution and abundance patterns. This study searches for community assembly rules by testing for coherence among functional groups and generalized guilds, as well as for correlations between the individual functional traits and assembly features, in two wetland plant communities in South Africa. The classifications of functional groups and generalized guilds were not consistent. However, rhizome internode length was related to fine-scale spatial pattern, suggesting that in systems dominated by clonal species (including wetlands, where recruitment sites are strongly limited) community assembly may be strongly linked to colonization ability. Functional groups do not predict guilds in wetland plant communities, precluding their use as the basis for assembly rules. However, an explicit consideration of clonal strategies and their effect on species’ spatial patterns appears to be important for understanding community assembly in systems dominated by clonal plants.     

Does post-fire abiotic habitat filtering create divergent plant communities in black spruce forests of eastern Canada?

We investigated the role of post-fire residual organic matter (ROM) thickness as a driver of community assembly in eastern Newfoundland. We hypothesized that if post-fire community assembly is predominantly controlled by ROM thickness (an abiotic habitat filter), then post-fire species composition and functional traits should correspond to the depth and distribution of ROM. However, if species interactions (biotic filter) are the primary constraints on community assembly, then post-fire species composition and their functional traits should be independent of the depth and distribution of ROM. We tested these predictions in three relatively mature plant communities, Kalmia angustifolia heath, black spruce (Picea mariana)-Kalmia shrub savannah and black spruce forest. Through pre-fire stand reconstruction, we found evidence that the three communities originated from black spruce forest. ROM thickness in heath was almost twice that of shrub savannah and six times more than forest, suggesting a gradient in fire severity. Distribution of ROM corresponded to patterns in vegetation dominance, where thick ROM (>2 cm) filtered out black spruce in favour of Kalmia. ROM thickness was a strong predictor of vegetation composition and function between heath and forest, but this was not found between the shrub savannah and forest. We attribute this to species interactions and allelopathy, which may have become important when ROM thickness was suitable for both seed (black spruce) and vegetative (Kalmia) regenerating species. Thus, priority effects or “who came first” may have lead to shrub savannah formation when ROM thickness was ~2 cm. We conclude that abiotic habitat filtering of thick ROM (>2 cm) on (primarily) species’ regeneration traits was the primary driver of community divergence from forest to heath and shrub savannah.     

Associations between floral specialization and species diversity: cause, effect, or correlation?

It has been proposed frequently, from Darwin’s time onwards, that specialized pollination increases speciation rates and thus the diversity of plant species (i.e. clade species richness). We suggest here that the correlation between clade species richness and floral specialization is real, but that clade species richness is frequently the cause, not the result of floral specialization. We urge a broader, variance-partitioning perspective for assessing the causes of this correlation by suggesting four models of how the diversity-specialization correlation might come about: (1) floral specialization promotes initial reproductive isolation (“Initial-RI” model), (2) floral specialization promotes reinforcement of reproductive isolation upon secondary contact (“Reinforcement” model), (3) floral specialization reduces the extinction rate by promoting tighter species packing (“Extinction” model), (4) floral specialization is the result of high clade species richness, which increases the number of related species in communities, and thus selects for floral character displacement (“Character-Displacement” model). These hypotheses are evaluated by comparing the relationships between species richness, speciation mechanisms, and pollination precision, accuracy, and specialization in the broader literature and, more specifically, in four study systems: Dalechampia (Euphorbiaceae), Collinsia (Plantaginaceae), Burmeistera (Campanulaceae), and Stylidium (Stylidiaceae). These systems provide stronger support for the character-displacement hypothesis, wherein local species diversity drives the evolution of specialized pollination. Although the two reproductive-isolation hypotheses may hold for plants like orchids, with extremely precise pollination systems, the reproductive character-displacement hypothesis seems likely to be more important for plant groups with less precise pollination systems.     

Measuring Functional Diversity in Plant Communities with MixedLife Forms: A Problem of Hardand Soft Attributes

This paper examines the functional relationships among species in an Australian rangeland community with mixed life forms. It follows a previous study (Walker and others 1999) that explored the role of dominant and minor species in maintaining functional diversity and resilience in a rangeland ecosystem. Unlike our previous results, which were based on estimates of five plant functional attributes, the dominant species in this second community apparently are functionally no more dissimilar to each other than to all other species. We suggest that the lack of clear results in mixed life form communities represents a confounding of the relationships between the “hard” attributes that actually govern how a plant performs in an ecosystem and the “soft” attributes that we use as surrogates. There are very few data on the hard functional attributes of plant species and consequently little information on precisely how the (soft) measurable traits are related to their imputed functions. What evidence there is shows that although the relationships are strong within life forms, they differ between life forms. This poses a problem for the development of research relating plant biodiversity to ecosystem function. Until such a database is developed, it will be very difficult to advance our understanding and measurement of functional diversity in mixed life form communities. Received 30 April 2001; Accepted 23 January 2002.     

Quantifying the role of deterministic assembly and stochastic drift in a natural community of Arctic mosses

The interpretation of natural plant communities frequently invokes species‐sorting controlled by niche differences along spatial environmental gradients. This process of niche structuring can be explained by reference to functional traits, which provide a mechanistic explanation for community structure. In contrast, models explaining species coexistence obviate the limiting effect of niche difference, by invoking processes which cause species‐level drift, e.g. demographic stochasticity. This paper investigates a simple habitat with strong gradients (moss communities in a patterned arctic wetland) to identify signature‐patterns under‐pinning the relative importance of deterministic assembly and stochastic drift in a natural community. First, ordination analysis was used to confirm community composition structured by a range of nine carefully selected functional traits. Second, to determine whether traits explaining community composition might also explain species richness, local species richness (sR) was compared to (1) observed trait diversity and (2) expected trait diversity based on permutation tests, which are used to simulate null community assembly for different values of sR. Traits explaining species composition, consistent with deterministic niche structuring, do not appear to maintain sR. This surprising result was explained by decomposing the community into individual pair‐wise comparisons, i.e. species niche‐differences and association (χ²). Results support deterministic processes via the sorting of species with similar and contrasting niches, at opposite ends of a composite environmental gradient. Nevertheless, stochastic drift is apparent in the random structure of a majority of pair‐wise associations; in addition, a species’ abundance was in general not related to environmental distance from response‐optima. We suggest therefore that spatial pattern in the moss community is a balance between deterministic forces with respect to species traits and controlling environmental gradients, and stochastic drift, which weakens this deterministic structure.     

Differences in functional trait responses to elevation among feeding guilds of Aculeata community

The response of communities to environmental change is expected to vary among feeding guilds. To evaluate the response of guilds to environmental factors without considering the taxonomic specificities, it is useful to examine Aculeata bees and wasps, which consist of closely related taxa including different guilds, pollinators, predators, and parasitoids. In this study, we evaluated changes in species diversity (SD) and functional traits of each feeding guild along an elevational gradient in a boreal forest in northern Japan. We used yellow pan traps to collect Aculeata bees and wasps at 200–1600 m above sea level. We investigated six functional traits (trophic level, seasonal duration, body size, elevational range, nesting position, and soil dependency) and the horizontal distribution of the species. The SD of all Aculeata, predators, and parasitoids decreased with an increase in elevation; however, the SD of pollinators did not show any specific trend. Although the functional trait composition of all Aculeata species did not show any trend, that of each feeding guild responded to elevation in different ways. Pollinators increased in body size and showed a decrease in seasonal duration with increasing elevation, suggesting that tolerance and seasonal escape from physical stress at high elevations are important for shaping pollinator communities. Predators increased their elevational range and the proportion of above‐ground nesting species increased with increasing elevation, suggesting that the ability to live in a wider range of environments and avoid unsuitable soil environments at high elevations might be important. Parasitoids changed their hosts and displayed variable traits with increasing elevation, suggesting that brood parasitoids have difficulty in surviving at high elevation. The traits for each guild responded in different ways, even if they were dominated by the same environmental factors. Our findings imply that differences in the responses of functional traits would produce different community assembly patterns in different guilds during further climate change.     

A predictive model of community assembly that incorporates intraspecific trait variation

Community assembly involves two antagonistic processes that select functional traits in opposite directions. Environmental filtering tends to increase the functional similarity of species within communities leading to trait convergence, whereas competition tends to limit the functional similarity of species within communities leading to trait divergence. Here, we introduce a new hierarchical Bayesian model that incorporates intraspecific trait variation into a predictive framework to unify classic coexistence theory and evolutionary biology with recent trait‐based approaches. Model predictions exhibited a significant positive correlation (r = 0.66) with observed relative abundances along a 10 °C gradient in mean annual temperature. The model predicted the correct dominant species in half of the plots, and accurately reproduced species' temperature optimums. The framework is generalizable to any ecosystem as it can accommodate any species pool, any set of functional traits and multiple environmental gradients, and it eliminates some of the criticisms associated with recent trait‐based community assembly models.     

Does Land-Use Intensification Decrease Plant Phylogenetic Diversity in Local Grasslands?

Phylogenetic diversity (PD) has been successfully used as a complement to classical measures of biological diversity such as species richness or functional diversity. By considering the phylogenetic history of species, PD broadly summarizes the trait space within a community. This covers amongst others complex physiological or biochemical traits that are often not considered in estimates of functional diversity, but may be important for the understanding of community assembly and the relationship between diversity and ecosystem functions. In this study we analyzed the relationship between PD of plant communities and land-use intensification in 150 local grassland plots in three regions in Germany. Specifically we asked whether PD decreases with land-use intensification and if so, whether the relationship is robust across different regions. Overall, we found that species richness decreased along land-use gradients the results however differed for common and rare species assemblages. PD only weakly decreased with increasing land-use intensity. The strength of the relationship thereby varied among regions and PD metrics used. From our results we suggest that there is no general relationship between PD and land-use intensification probably due to lack of phylogenetic conservatism in land-use sensitive traits. Nevertheless, we suggest that depending on specific regional idiosyncrasies the consideration of PD as a complement to other measures of diversity can be useful.     

Eco-evolutionary dynamics in herbivorous insect communities mediated by induced plant responses

It is increasingly recognized that the ecology of communities and evolution of species within communities are interdependent, and researchers have been paying attention to this rapidly emerging field of research, i.e., through studies on eco-evolutionary dynamics. Most of the studies on eco-evolutionary dynamics have been concerned with direct trophic interactions. However, community ecologists have shown that trait-mediated indirect effects play an important role in shaping the structure of natural communities. In particular, in terrestrial plant–insect systems, indirect effects mediated through herbivore-induced plant responses are common and have a great impact on the structure of herbivore communities. This review describes eco-evolutionary dynamics in herbivorous insect communities, and specifically focuses on the key role of herbivore-induced plant responses in eco-evolutionary dynamics. First, I review studies on the evolution of herbivore traits relevant to plant induction and discuss evolution in a community context mediated by induced plant responses. Second, I highlight how intraspecific genetic variation or evolution in herbivore traits can influence herbivore community structure. Finally, I propose the hypothetical model that induced plant responses supports eco-evolutionary feedback in herbivore communities. In this review, I argue that the application of the indirect interaction web approaches into studies on eco-evolutionary will provide profound insights into understanding of mechanisms of the generation and maintenance of biodiversity.     

Isolation‐driven functional assembly of plant communities on islands

The physical and biotic environment is often considered the primary driver of functional variation in plant communities. Here, we examine the hypothesis that spatial isolation may also be an important driver of functional variation in plant communities where disturbance and dispersal limitation may prevent species from occupying all suitable habitats. To test this hypothesis, we surveyed the vascular plant composition of 30 islands in the Gulf of Maine, USA, and used available functional trait and growth form data to quantify the functional composition of these islands. We categorized species based on dispersal mode and used a landscape metric of isolation to assess the potential role of dispersal limitation as a mechanism of isolation‐driven assembly. We tested for island and species level effects on functional composition using a hierarchical Bayesian framework to better assess the causal link between isolation and functional variation. Growth form composition and the community mean value of functional traits related to growth rate, stress tolerance, and nutrient use varied significantly with island isolation. Functional traits and growth forms were significantly associated with dispersal mode, and spatial isolation was the strongest driver of primary trait variation, while island properties associated with environmental drivers in our system were not strong predictors of trait variation. Despite the species‐level association of dispersal mode and functional traits, dispersal mode only accounted for a small proportion of the overall isolation effect on community‐level trait variation. Our study suggests that spatial isolation can be a key driver of functional assembly in plant communities on islands, though the role of particular dispersal processes remains unclear.