Dissecting spatiotemporal patterns of functional diversity through the lens of Darwin's naturalization conundrum

Darwin's naturalization conundrum describes the paradigm that community assembly is regulated by two opposing processes, environmental filtering and competitive interactions, which predict both similarity and distinctiveness of species to be important for establishment. Our goal is to use long‐term, large‐scale, and high‐resolution temporal data to examine diversity patterns over time and assess whether environmental filtering or competition plays a larger role in regulating community assembly processes. We evaluated Darwin's naturalization conundrum and how functional diversity has changed in the Laurentian Great Lakes fish community from 1870 to 2010, which has experienced frequent introductions of non‐native species and extirpations of native species. We analyzed how functional diversity has changed over time by decade from 1870 to 2010 at three spatial scales (regional, lake, and habitat) to account for potential noninteractions between species at the regional and lake level. We also determined which process, environmental filtering or competitive interactions, is more important in regulating community assembly and maintenance by comparing observed patterns to what we would expect in the absence of an ecological mechanism. With the exception of one community, all analyses show that functional diversity and species richness has increased over time and that environmental filtering regulates community assembly at the regional level. When examining functional diversity at the lake and habitat level, the regulating processes become more context dependent. This study is the first to examine diversity patterns and Darwin's conundrum by integrating long‐term, large‐scale, and high‐resolution temporal data at multiple spatial scales. Our results confirm that Darwin's conundrum is highly context dependent.     

Advances, challenges and a developing synthesis of ecological community assembly theory

Ecological approaches to community assembly have emphasized the interplay between neutral processes, niche-based environmental filtering and niche-based species sorting in an interactive milieu. Recently, progress has been made in terms of aligning our vocabulary with conceptual advances, assessing how trait-based community functional parameters differ from neutral expectation and assessing how traits vary along environmental gradients. Experiments have confirmed the influence of these processes on assembly and have addressed the role of dispersal in shaping local assemblages. Community phylogenetics has forged common ground between ecologists and biogeographers, but it is not a proxy for trait-based approaches. Community assembly theory is in need of a comparative synthesis that addresses how the relative importance of niche and neutral processes varies among taxa, along environmental gradients, and across scales. Towards that goal, we suggest a set of traits that probably confer increasing community neutrality and regionality and review the influences of stress, disturbance and scale on the importance of niche assembly. We advocate increasing the complexity of experiments in order to assess the relative importance of multiple processes. As an example, we provide evidence that dispersal, niche processes and trait interdependencies have about equal influence on trait-based assembly in an experimental grassland.     

Contribution of non-native species to soft-sediment marine community structure of San Francisco Bay,California

While non-native species (NIS) are important components in many coastal bays and estuaries, quantitative measures that characterize their effects on community structure at bay-wide scales are rare. In this study, we measure species composition and abundance for soft-sediments to assess the contribution of NIS to multiple dimensions of community structure, focusing on one of the most highly invaded bays in the world, San Francisco Bay. Benthic macrofauna was sampled in the high salinity, muddy shallow subtidal (2 m depth) across 10 sites, using replicate 0.1 m² Van Veen grabs. Invertebrates retained on a 1 mm sieve were identified, counted, and used to estimate the overall contribution of NIS to (a) abundance (b) species richness, and (c) community similarity. Soft-sediment communities were dominated numerically by NIS, which accounted for 76 % of all organisms detected and had a mean bay-wide abundance that was three and half-fold higher than native biota. Overall, NIS contributed to 36 % of observed taxa and 24–29 % of total estimated regional diversity. Native species accounted for 21 % of total abundance and 45 % of total species richness. Compared to native species, NIS occurred more frequently among samples and also explained more of the variation in community structure among sites. NIS dominate several key attributes of the soft-sediment infaunal community in San Francisco Bay. Percent contribution of NIS to species richness was at least two-fold higher than reported from two decades ago. Unique to this bay, these measures establish a quantitative baseline on the state of invasions and provide an important model for evaluating the extent of NIS in estuaries. Application of this approach across estuaries, with repeated measures over time, is critically needed to advance scientific understanding of invasions and also evaluation of efficacy and gaps in management to reduce new invasions.     

Phylogenetic community ecology and the role of social dominance in sponge-dwelling shrimp

When functional traits are evolutionarily conserved, phylogenetic relatedness can serve as a proxy for ecological similarity to examine whether functional differences among species mediate community assembly. Using phylogenetic- and trait-based analyses, we demonstrate that sponge-dwelling shrimp (Synalpheus) assemblages are structured by size-based habitat filtering, interacting with competitive exclusion mediated by social system. Most shrimp communities were more closely related and/or more similar in size than randomized communities, consistent with habitat filtering facilitated by phylogenetically conserved body size. Those sponges with greater space heterogeneity hosted shrimp communities with greater size diversity, corroborating the importance of size in niche use. However, communities containing eusocial shrimp - which cooperatively defend territories - were less phylogenetically related and less similar in size, suggesting that eusociality enhances competitive ability and drives competitive exclusion. Our analyses demonstrate that community assembly in this diverse system occurs via traits mediating niche use and differential competitive ability.     

Distribution of functionally distinct native and non-indigenous species within marine urban habitats

Aim

Trait-based approaches are powerful to examine the processes associated with biological invasions. Functional comparison among native and non-indigenous species (NIS) can notably infer whether novel assemblages result from neutral or niche-based assembly rules. Applying such a framework to biofouling communities, our study aimed to elucidate their distributions within two marine urban habitats (namely floating vs. nonfloating habitats).

Location

Southeast Pacific—Central Chilean coastline.

Methods

Here, we examined the distribution of 12 functional traits in fouling communities established on settlement plates, after 3 and 13 months of deployment in the two habitats and across ports in Central Chile. Based upon previously described differences of assemblages and NIS contribution across habitats, we hypothesized that nonindigenous, cryptogenic and native taxon pools would be functionally distinct (and trait biased), and that functional diversity and structure would vary across habitats and successional stages.

Results

Our results show, as anticipated, that nonindigenous (13 taxa), cryptogenic (12) and native (18) taxon pools are functionally distinct, though overlapping in the trait space. Non-indigenous species are rather related to colonizing traits, while native species are more related to competitive traits. Only one widespread NIS was functionally similar to the late successional and most competitive native species, including taxa elsewhere invasives. Despite differences in taxonomic composition between habitats, we did not observe functional differences between them. In contrast, temporal variations across colonization stages were detected along with an increased contribution in large and long-lived taxa, together with site-specific trajectories.

Main Conclusions

We conclude that the functional distinctness among nonindigenous, cryptogenic and native taxa occupying artificial habitats in ports reflects niche-based processes. Site-specific trajectories indicate that scale-dependent assembly processes, such as dispersal and species interactions, are at play.     

The Influence of Matrix Size on Statistical Properties of Co-Occurrence and Limiting Similarity Null Models

Null models exploring species co-occurrence and trait-based limiting similarity are increasingly used to explore the influence of competition on community assembly; however, assessments of common models have not thoroughly explored the influence of variation in matrix size on error rates, in spite of the fact that studies have explored community matrices that vary considerably in size. To determine how smaller matrices, which are of greatest concern, perform statistically, we generated biologically realistic presence-absence matrices ranging in size from 3–50 species and sites, as well as associated trait matrices. We examined co-occurrence tests using the C-Score statistic and independent swap algorithm. For trait-based limiting similarity null models, we used the mean nearest neighbour trait distance (NN) and the standard deviation of nearest neighbour distances (SDNN) as test statistics, and considered two common randomization algorithms: abundance independent trait shuffling (AITS), and abundance weighted trait shuffling (AWTS). Matrices as small as three × three resulted in acceptable type I error rates (p < 0.05) for both the co-occurrence and trait-based limiting similarity null models when exclusive p-values were used. The commonly used inclusive p-value (≤ or ≥, as opposed to exclusive p-values; < or >) was associated with increased type I error rates, particularly for matrices with fewer than eight species. Type I error rates increased for limiting similarity tests using the AWTS randomization scheme when community matrices contained more than 35 sites; a similar randomization used in null models of phylogenetic dispersion has previously been viewed as robust. Notwithstanding other potential deficiencies related to the use of small matrices to represent communities, the application of both classes of null model should be restricted to matrices with 10 or more species to avoid the possibility of type II errors. Additionally, researchers should restrict the use of the AWTS randomization to matrices with fewer than 35 sites to avoid type I errors when testing for trait-based limiting similarity. The AITS randomization scheme performed better in terms of type I error rates, and therefore may be more appropriate when considering systems for which traits are not clustered by abundance.     

Fish species introductions provide novel insights into the patterns and drivers of phylogenetic structure in freshwaters

Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. We quantify phylogenetic clustering and overdispersion of native and non-native fishes of a large river basin in the American Southwest to test for the mechanisms (environmental filtering versus competitive exclusion) and spatial scales influencing community structure. Contrary to expectations, non-native species were phylogenetically clustered and related to natural environmental conditions, whereas native species were not phylogenetically structured, likely reflecting human-related changes to the basin. The species that are most invasive (in terms of ecological impacts) tended to be the most phylogenetically divergent from natives across watersheds, but not within watersheds, supporting the hypothesis that Darwin''s naturalization conundrum is driven by the spatial scale. Phylogenetic distinctiveness may facilitate non-native establishment at regional scales, but environmental filtering restricts local membership to closely related species with physiological tolerances for current environments. By contrast, native species may have been phylogenetically clustered in historical times, but species loss from contemporary populations by anthropogenic activities has likely shaped the phylogenetic signal. Our study implies that fundamental mechanisms of community assembly have changed, with fundamental consequences for the biogeography of both native and non-native species.     

Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits??vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)??in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.     

A trait-based approach predicting community assembly and dominance of microbial invasive species

Understanding the mechanisms underlying community assembly helps to define success and susceptibility to biological invasions. Here, we explored phytoplankton community assembly following niche and neutral paradigms and using a trait-based approach. Under the hypothesis that the morphology-based functional groups (MBFG) clusters species with similar niche, we analysed how trait-related differences in fitness influence dominance of an invasive species. This was based on literature review, field data and model simulations. We predict that invading species can be dominant if: 1) do not belong to the local MBFG but use unexploited areas of the niche, or 2) belong to the resident MBFG but exhibit a higher fitness due to a particular combination of traits. The invasive dinoflagellate Ceratium furcoides was used as the model species to evaluate these hypotheses, its morphological (e.g. volume) and physiological (e.g. growth rates) traits were compared with species from the same (V: photosynthetic flagellates) and different (VII: colonial cyanobacteria) MBFG. Fitness was estimated using models parametrized with MBFG rates (R*, ability to draw down phosphate) under different environmental conditions (i.e. flushing). Results contributed to support both hypotheses. First, the alternation of C. furcoides and cyanobacteria dominance was explained by the use of different niches. Secondly, species from MBFG V were dominant under similar environments. Within this group V C. furcoides showed higher fitness under low flushing and high predation, advantage provided by a distinctive combination of traits. The application of trait-based approaches to represent the niche and estimate fitness along environmental gradients was useful to evaluate community assembly and can be used to predict the dominance of microbial species invasions.     

Non-Native Plants Disrupt Dual Promotion of Native Alpha and Beta Diversity

Non-native species can alter patterns of species diversity at multiple spatial scales, but the processes that underlie multi-scale effects remain unclear. Here we show that non-native species reduce native diversity at multiple scales through simultaneous disruption of two processes of native community assembly: species immigration, which enhances alpha diversity, and community divergence, which enhances beta diversity. Community divergence refers to the process in which local communities diverge over time in species composition because the history of species immigration and, consequently, the way species affect one another within communities are variable among communities. Continuous experimental removal of species over four years of floodplain succession revealed that, when non-native species were excluded, stochastic variation in the timing of a dominant native species’ arrival allowed local communities to diverge, thereby enhancing beta diversity, without compromising promotion of alpha diversity by species immigration. In contrast, when non-native species were allowed to enter experimental plots, they not only reduced native alpha diversity by limiting immigration, but also diminished the dominant native species’ role in enhancing native beta diversity. Our results highlight the importance of community assembly and succession for understanding multi-scale effects of non-native species.     

Using Plant Functional Traits and Phylogenies to Understand Patterns of Plant Community Assembly in a Seasonal Tropical Forest in Lao PDR

Plant functional traits reflect different evolutionary responses to environmental variation, and among extant species determine the outcomes of interactions between plants and their environment, including other plant species. Thus, combining phylogenetic and trait-based information can be a powerful approach for understanding community assembly processes across a range of spatial scales. We used this approach to investigate tree community composition at Phou Khao Khouay National Park (18°14’-18°32’N; 102°38’- 102°59’E), Laos, where several distinct forest types occur in close proximity. The aim of our study was to examine patterns of plant community assembly across the strong environmental gradients evident at our site. We hypothesized that differences in tree community composition were being driven by an underlying gradient in soil conditions. Thus, we predicted that environmental filtering would predominate at the site and that the filtering would be strongest on sandier soil with low pH, as these are the conditions least favorable to plant growth. We surveyed eleven 0.25 ha (50x50 m) plots for all trees above 10 cm dbh (1221 individual trees, including 47 families, 70 genera and 123 species) and sampled soils in each plot. For each species in the community, we measured 11 commonly studied plant functional traits covering both the leaf and wood economic spectrum traits and we reconstructed a phylogenetic tree for 115 of the species in the community using rbcL and matK sequences downloaded from Genebank (other species were not available). Finally we compared the distribution of trait values and species at two scales (among plots and 10x10m subplots) to examine trait and phylogenetic community structures. Although there was strong evidence that an underlying soil gradient was determining patterns of species composition at the site, our results did not support the hypothesis that the environmental filtering dominated community assembly processes. For the measured plant functional traits there was no consistent pattern of trait dispersion across the site, either when traits were considered individually or when combined in a multivariate analysis. However, there was a significant correlation between the degree of phylogenetic dispersion and the first principle component axis (PCA1) for the soil parameters. Moreover, the more phylogenetically clustered plots were on sandier soils with lower pH. Hence, we suggest that the community assembly processes across our site may reflect the influence of more conserved traits that we did not measure. Nevertheless, our results are equivocal and other interpretations are possible. Our study illustrates some difficulties in combining trait and phylogenetic approaches that may result from the complexities of integrating spatial and evolutionary processes that vary at different scales.     

Are urban systems beneficial,detrimental, or indifferent for biological invasion?

Urban environments are often seen as unique or degraded habitats that both present hardships for some sensitive species and provide opportunities to others. Non-indigenous species (NIS) are commonly referenced in the latter group, and are comprised of species that can tolerate the unique conditions or capitalize on the opportunities found in urban environments. Moreover, these urban beneficiaries may be those that normally cannot overcome competitive interactions in intact native communities, but find opportunity to flourish in urban habitats. We ask the question: do NIS benefit from urbanization? We answer this question using three strategies. First, we explore the problem conceptually, using community assembly theory. Second, we perform a broad literature review. Finally, we analyze studies with sufficient information using a meta-analysis. We show that the available evidence supports the proposition that NIS benefit from urbanization, with NIS obtaining higher abundances and greater diversity in more urbanized habitats. There were only 43 studies that measured NIS abundance and diversity while adequately quantifying the degree of urbanization surrounding plots, and effect sizes (measured by Hedge’s D) reveal that NIS obtain higher abundances in more urbanized habitats, and especially for invertebrates. Despite the intense interest in NIS dynamics and impacts, we note a general dearth of robust studies that adequately quantify ‘urbanization’, and we end with a general call for more detailed research.     

Changing assembly rules during secondary succession: evidence for non-random patterns

Describing the rules of community assembly is a central topic of ecology. Studying successional processes through a trait-based null model approach can help to better understand the rules of community assembly.According to theoretical considerations, at the beginning of succession - after getting over the dispersal limitation stage - community composition is primarily shaped by environmental filters (generating functional convergence), while in later stages limiting similarity (generating functional divergence) will be dominant. However, empirical evidence does not clearly support theoretical expectations.Our aim was to detect the presence and changes of trait-based assembly processes during old-field succession based on twelve traits. Changes in vegetation composition were evaluated by a combination of time series and space-for-time substitution: conducting three resurveys of permanent plots on four old-field age-groups. The individual dispersion of traits was transformed into effect size (i.e. departure from null model expectation). The impact of time since abandonment on effect sizes was tested by generalized additive mixed effect models.We detected a non-random pattern for each trait in at least some part of the succession. Departure from randomness did not change significantly over time for six traits: seed mass, lateral spread and pollination type were divergent, while leaf size, generative height and length of flowering were convergent. Six traits had changing patterns along the succession. Four of them showed increasing divergence (e.g. dispersal type, LDMC), which supports our hypothesis. While two (SLA, life form) displayed increasing convergence, contrary to expectations.We confirmed the general hypothesis that convergence is predominant initially and that divergence can be detected later in succession for four traits. However, the large variation found in trait dispersion indicates that complex processes operate during succession.     

Is invasion success explained by the enemy release hypothesis?

A recent trend in invasion ecology relates the success of non‐indigenous species (NIS) to reduced control by enemies such as pathogens, parasites and predators (i.e. the enemy release hypothesis, ERH). Despite the demonstrated importance of enemies to host population dynamics, studies of the ERH are split – biogeographical analyses primarily show a reduction in the diversity of enemies in the introduced range compared with the native range, while community studies imply that NIS are no less affected by enemies than native species in the invaded community. A broad review of the invasion literature implies at least eight non‐exclusive explanations for this enigma. In addition, we argue that the ERH has often been accepted uncritically wherever (i) NIS often appear larger, more fecund, or somehow ‘better’ than either congeners in the introduced region, or conspecifics in the native range; and (ii) known enemies are conspicuously absent from the introduced range. However, all NIS, regardless of their abundance or impact, will lose natural enemies at a biogeographical scale. Given the complexity of processes that underlie biological invasions, we argue against a simple relationship between enemy ‘release’ and the vigour, abundance or impact of NIS.     

Land‐use and isolation interact to affect wetland plant assemblages

Different management regimes imposed on similar habitat types provide opportunities to investigate mechanisms driving community assembly and changes in species composition. We investigated the effect of pasture management on vegetation composition in wetlands with varying spatial isolation on a Florida cattle ranch. We hypothesized that increased pasture management intensity would dampen the expected negative effect of wetland isolation on native species richness due to a change from dispersal‐driven community assembly to niche‐driven assembly by accentuated environmental tolerance. We used native plant richness, exotic plant richness and mean coefficient of conservatism (CC) to assess wetland plant assemblage composition. Sixty wetlands were sampled, stratified by three levels of isolation across two pasture management intensities; semi‐native (less intensely managed; mostly native grasses, never fertilized) and agronomically improved (intensely managed, planted with exotic grasses, and fertilized). Improved pasture wetlands had lower native richness and CC scores, and greater total soil phosphorus and exotic species coverage compared to semi‐native pasture wetlands. Increased wetland isolation was significantly associated with decreases in native species richness in semi‐native pasture wetlands but not in improved pasture wetlands. Additionally, the species–area relationship was stronger in wetlands in improved pastures than semi‐native pastures. Our results indicate that a) native species switch from dispersal‐based community assembly in semi‐native pastures to a species‐sorting process in improved pastures, and b) recently‐introduced exotic species already sorted for more intensive management conditions are primarily undergoing dispersal‐based community assembly. That land‐use may alter the relative importance of assembly processes and that different processes drive native and exotic richness has implications for both ecosystem management and restoration planning.     

The assembly of tropical tree communities – the advances and shortcomings of phylogenetic and functional trait analyses

Tropical tree communities present one of the most challenging systems for studying the processes underlying community assembly. Most community assembly hypotheses consider the relative importance of the ecological similarity of co‐occurring species. Quantifying this similarity is a daunting and potentially impossible task in species‐rich assemblages. During the past decade tropical tree ecologists have increasingly utilized phylogenetic trees and functional traits to estimate the ecological similarity of species in order to test mechanistic community assembly hypotheses. A large amount of work has resulted with many important advances having been made along the way. That said, there are still many outstanding challenges facing those utilizing phylogenetic and functional trait approaches to study community assembly. Here I review the conceptual background, major advances and major remaining challenges in phylogenetic‐ and trait‐based approaches to community ecology with a specific focus on tropical trees. I argue that both approaches tremendously improve our understanding of tropical tree community ecology, but neither approach has fully reached its potential thus far.     

Do pollinators influence the assembly of flower colours within plant communities?

The co-occurrence of plant species within a community is influenced by local deterministic or neutral processes as well as historical regional processes. Floral trait distributions of co-flowering species that share pollinators may reflect the impact of pollinator preference and constancy on their assembly within local communities. While pollinator sharing may lead to increased visitation rates for species with similar flowers, the receipt of foreign pollen via interspecific pollinator movements can decrease seed set. We investigated the pattern of community flower colour assembly as perceived by native honeybee pollinators within 24 local assemblages of co-flowering Oxalis species within the Greater Cape Floristic Region, South Africa. To explore the influence of pollinators on trait assembly, we assessed the impact of colour similarity on pollinator choices and the cost of heterospecific pollen receipt. We show that flower colour is significantly clustered within Oxalis communities and that this is not due to historical constraint, as flower colour is evolutionarily labile within Oxalis and communities are randomly structured with respect to phylogeny. Pollinator observations reveal that the likelihood of pollinators switching between co-flowering species is low and increases with flower colour similarity. Interspecific hand pollination significantly reduced seed set in the four Oxalis species we investigated, and all were dependant on pollinators for reproduction. Together these results imply that flower colour similarity carries a potential fitness cost. However, pollinators were highly flower constant, and remained so despite the extreme similarity of flower colour as perceived by honeybees. This suggests that other floral traits facilitate discrimination between similarly coloured species, thereby likely resulting in a low incidence of interspecific pollen transfer (IPT). If colour similarity promotes pollinator attraction at the community level, the observed clustering of flower colour within communities might result from indirect facilitative interactions.     

Pollution reduces native diversity and increases invader dominance in marine hard-substrate communities

Anthropogenic disturbance is considered a risk factor in the establishment of non‐indigenous species (NIS); however, few studies have investigated the role of anthropogenic disturbance in facilitating the establishment and spread of NIS in marine environments. A baseline survey of native and NIS was undertaken in conjunction with a manipulative experiment to determine the effect that heavy metal pollution had on the diversity and invasibility of marine hard‐substrate assemblages. The study was repeated at two sites in each of two harbours in New South Wales, Australia. The survey sampled a total of 47 sessile invertebrate taxa, of which 15 (32%) were identified as native, 19 (40%) as NIS, and 13 (28%) as cryptogenic. Increasing pollution exposure decreased native species diversity at all study sites by between 33% and 50%. In contrast, there was no significant change in the numbers of NIS. Percentage cover was used as a measure of spatial dominance, with increased pollution exposure leading to increased NIS dominance across all sites. At three of the four study sites, assemblages that had previously been dominated by natives changed to become either extensively dominated by NIS or equally occupied by native and NIS alike. No single native or NIS was repeatedly responsible for the observed changes in native species diversity or NIS dominance at all sites. Rather, the observed effects of pollution were driven by a diverse range of taxa and species. These findings have important implications for both the way we assess pollution impacts, and for the management of NIS. When monitoring the response of assemblages to pollution, it is not sufficient to simply assess changes in community diversity. Rather, it is important to distinguish native from NIS components since both are expected to respond differently. In order to successfully manage current NIS, we first need to address levels of pollution within recipient systems in an effort to bolster the resilience of native communities to invasion.     

Contrasting patterns of trait‐based community assembly in lianas and trees from temperate Australia

Trait variation in plant communities is thought to be constrained by two opposing community assembly processes operating at discrete spatial scales: habitat filtering and limiting similarity between coexisting species. Filtering processes cause convergence in ecological strategy as species are excluded from unsuitable sites, whilst limiting similarity leads to the divergence of trait values between co‐occurring species in order to alleviate competition for finite resources. Levels of alpha (within‐site) and beta (among‐site) trait variation can be indicative of the strength of these community assembly processes. We used trait‐gradient analysis to explicitly compare evidence of community assembly patterns in lianas (woody vines) and trees. These two growth forms exhibit striking differences in carbon capture and regeneration strategies, yet trait‐based mechanisms that maintain their coexistence remain understudied. Using data for four functional traits – leaf mass per area, leaf nitrogen content (N_mass), leaf area and seed mass – we partitioned interspecific trait variation in lianas and trees into alpha and beta components. Our three key findings were: 1) lianas and trees exhibit divergent patterns of trait‐based habitat filtering, due to differences in the relationship between leaf size and the other three traits examined (LMA, N_mass and seed mass), 2) on average, liana species possess smaller seeds, lower LMA and higher N_mass than do trees, but there was no clear difference in leaf area between the two growth forms, and 3) soil fertility was correlated with trait variation (leaf area, seed mass) at the site‐level in trees, but not in lianas. These results provide evidence that dominant growth forms can be filtered into the same habitat on the basis of different combinations of traits. Our findings have important implications for community assembly and co‐existence theory and for more pragmatic matters such as using trait‐based principles to inform community restoration.     

A trait-based approach across the native and invaded range to understand plant invasiveness and community impact

Alien plant species invasiveness and impact on diversity (i.e. species richness and composition) can be driven by the altered competitive interactions experienced by the invader in its invaded range compared to its native range. Trait-based competition effects on invasiveness can be mediated through size-asymmetric competition, i.e. a trait suit of the invader that drives competitive dominance, and through ‘niche differences', i.e. trait differentiation and thus minimized competition between invader and the invaded community. In terms of invasion impact, size-asymmetric competition is expected to result in competitive exclusion of co-occurring subordinate species, whereas ‘niche differences' might result in competitive exclusion of the most functionally similar co-occurring species. Although observational work does not allow the full disentanglement of both trait-based effects, it does allow to verify the occurrence of expected theoretical trait patters. In this study, we explored the trait-based competition effects on invasiveness and diversity impact for Rosa rugosa in both its invaded range in Belgium and its native range in Japan, based on seven functional traits across 100 vegetation plots. Following the predictions for enhanced invasiveness, we found much lower functional overlap between R. rugosa and the co-occurring species in the invaded range compared to the native range. This likely also explains the absence of diversity impact in its native range. Despite the absence of changes in species richness in the invaded range, the invader did strongly impact species composition of invaded communities. This impact occurred through strong shade tolerance responses, suggesting size-asymmetric competition effects and cover loss of co-occurring dominant species, next to exclusion of co-occurring species most functionally similar to the invader; suggesting niche difference effects. In conclusion, this case-study illustrates how exploring functional trait patterns across a species native and invaded range can help in understanding how trait-based competition processes can affect invasiveness and community impact.