Turnover of plant lineages shapes herbivore phylogenetic beta diversity along ecological gradients

Understanding drivers of biodiversity patterns is of prime importance in this era of severe environmental crisis. More diverse plant communities have been postulated to represent a larger functional trait‐space, more likely to sustain a diverse assembly of herbivore species. Here, we expand this hypothesis to integrate environmental, functional and phylogenetic variation of plant communities as factors explaining the diversity of lepidopteran assemblages along elevation gradients in the Swiss Western Alps. According to expectations, we found that the association between butterflies and their host plants is highly phylogenetically structured. Multiple regression analyses showed the combined effect of climate, functional traits and phylogenetic diversity in structuring butterfly communities. Furthermore, we provide the first evidence that plant phylogenetic beta diversity is the major driver explaining butterfly phylogenetic beta diversity. Along ecological gradients, the bottom up control of herbivore diversity is thus driven by phylogenetically structured turnover of plant traits as well as environmental variables.     

Differential bacterial dynamics promote emergent community robustness to lake mixing: an epilimnion to hypolimnion transplant experiment

Lake mixing disrupts chemical and physical gradients that structure bacterial communities. A transplant experiment was designed to investigate the influence of post‐mixing environmental conditions and biotic interactions on bacterial community composition. The experimental design was 3 × 2 factorial, where water was incubated from three different sources (epilimnion, hypolimnion, and mixed epilimnion and hypolimnion) at two different locations in the water column (epilimnion or hypolimnion). Three replicate mesocosms of each treatment were removed every day for 5 days for bacterial community profiling, assessed by automated ribosomal intergenic spacer analysis. There were significant treatment effects observed, and temperature was the strongest measured driver of community change (r = ?0.66). Epilimnion‐incubated communities changed more than hypolimnion‐incubated. Across all treatments, we classified generalist, layer‐preferential and layer‐specialist populations based on occurrence patterns. Most classified populations were generalists that occurred in both strata, suggesting that communities were robust to mixing. In a network analysis of the mixed‐inocula treatments, there was correlative evidence of inter‐population biotic interactions, where many of these interactions involved generalists. These results reveal differential responses of bacterial populations to lake mixing and highlight the role of generalist taxa in structuring an emergent community‐level response.     

The influence of habitat heterogeneity on freshwater bacterial community composition and dynamics

Multiple forces structure natural microbial communities, but the relative roles and interactions of these drivers are poorly understood. Gradients of physical and chemical parameters can be especially influential. In traditional ecological theory, variability in environmental conditions across space and time represents habitat heterogeneity, which may shape communities. Here we used aquatic microbial communities as a model to investigate the relationship between habitat heterogeneity and community composition and dynamics. We defined spatial habitat heterogeneity as vertical temperature and dissolved oxygen (DO) gradients in the water column, and temporal habitat heterogeneity as variation throughout the open-water season in these environmental parameters. Seasonal lake mixing events contribute to temporal habitat heterogeneity by destroying and re-creating these gradients. Because of this, we selected three lakes along a range of annual mixing frequency (polymictic, dimictic, meromictic) for our study. We found that bacterial community composition (BCC) was distinct between the epilimnion and hypolimnion within stratified lakes, and also more variable within the epilimnia through time. We found stark differences in patterns of epilimnion and hypolimnion dynamics over time and across lakes, suggesting that specific drivers have distinct relative importance for each community.     

Phylogenetic structure and phylogenetic diversity of angiosperm assemblages in forests along an elevational gradient in Changbaishan,China

Aims Understanding what drives the variation in species composition and diversity among local communities can provide insights into the mechanisms of community assembly. Because ecological traits are often thought to be phylogenetically conserved, there should be patterns in phylogenetic structure and phylogenetic diversity in local communities along ecological gradients. We investigate potential patterns in angiosperm assemblages along an elevational gradient with a steep ecological gradient in Changbaishan, China.Methods We used 13 angiosperm assemblages in forest plots (32×32 m) distributed along an elevational gradient from 720 to 1900 m above sea level. We used Faith's phylogenetic diversity metric to quantify the phylogenetic alpha diversity of each forest plot, used the net relatedness index to quantify the degree of phylogenetic relatedness among angiosperm species within each forest plot and used a phylogenetic dissimilarity index to quantify phylogenetic beta diversity among forest plots. We related the measures of phylogenetic structure and phylogenetic diversity to environmental (climatic and edaphic) factors.Important findings Our study showed that angiosperm assemblages tended to be more phylogenetically clustered at higher elevations in Changbaishan. This finding is consistent with the prediction of the phylogenetic niche conservatism hypothesis, which highlights the role of niche constraints in governing the phylogenetic structure of assemblages. Our study also showed that woody assemblages differ from herbaceous assemblages in several major aspects. First, phylogenetic clustering dominated in woody assemblages, whereas phylogenetic overdispersion dominated in herbaceous assemblages; second, patterns in phylogenetic relatedness along the elevational and temperature gradients of Changbaishan were stronger for woody assemblages than for herbaceous assemblages; third, environmental variables explained much more variations in phylogenetic relatedness, phylogenetic alpha diversity and phylogenetic beta diversity for woody assemblages than for herbaceous assemblages.     

The building of a biodiversity hotspot across a land-bridge in the Mediterranean

Many of the macroevolutionary processes that have shaped present-day phylogenetic patterns were caused by geological events such as plate tectonics and temporary land-bridges. The study of spatial patterns of phylogenetic diversity can provide insights into these past events. Here we focus on a western Mediterranean biodiversity hotspot located in the southern Iberian Peninsula and northwest Africa, two regions that are separated by the Strait of Gibraltar. We explore the spatial structure of the phylogenetic relationships within and across large-scale plant assemblages. Significant turnover in terminal lineages tends to occur between landmasses, whereas turnover in deep lineages tends to occur within landmasses. Plant assemblages in the western ecoregions of this hotspot tend to be phylogenetically overdispersed but are phylogenetically clustered on its eastern margins. We discuss our results in the light of potential scenarios of niche evolution (or conservatism) and lineage diversification. The significant turnover between landmasses suggests a common scenario of allopatric speciation that could have been facilitated by the intermittent joining of the two continents. This may have constituted an important stimulus for diversification and the emergence of this western Mediterranean biodiversity hotspot.     

Global patterns in the phylogenetic structure of island mammal assemblages

Assemblage-level phylogenies carry the signature of ecological and evolutionary processes, which may provide useful information on modes of assemblage formation. We present a global-scale analysis of the emergent phylogenetic properties of mammal assemblages on islands, in which we compared the structure of 595 island assemblages with null models constructed under four alternative definitions of regional source pools. Although most assemblages had a structure indistinguishable from random samples, for some mammal taxa, up to 40% of island assemblages were phylogenetically overdispersed. This suggests that in at least some cases, the processes that shape island faunas are not independent of phylogeny. Furthermore, measures of phylogenetic structure were associated in some cases with island geographical features (size, maximum elevation and habitat diversity). Our results suggest that part of the signal of assemblage formation processes is detectable in the phylogenies of contemporary island mammal faunas, though much is obscured by the complexity of these processes.     

Phylogenetic structure of mammal assemblages at large geographical scales: linking phylogenetic community ecology with macroecology

Phylogenetic community ecology seeks to explain the processes involved in the formation of species assemblages by analysing their phylogenetic structure, and to date has focused primarily on local-scale communities. Macroecology, on the other hand, is concerned with the structure of assemblages at large geographical scales, but has remained largely non-phylogenetic. Analysing the phylogenetic structure of large-scale assemblages provides a link between these two research programmes. In this paper, I ask whether we should expect large-scale assemblages to show significant phylogenetic structure, by outlining some of the ecological and macroevolutionary processes that may play a role in assemblage formation. As a case study, I then explore the phylogenetic structure of carnivore assemblages within the terrestrial ecoregions of Africa. Many assemblages at these scales are indeed phylogenetically non-random (either clustered or overdispersed). One interpretation of the observed patterns of phylogenetic structure is that many clades underwent rapid biome-filling radiations, followed by diversification slowdown and competitive sorting as niche space became saturated.     

系统发育多样性与系统发育结构在岛屿植物群落保护中的意义——以蜈支洲岛为例

为探讨系统发育多样性和系统发育结构在岛屿植物群落保护决策制定中的作用,以海南三亚蜈支洲岛被子植物群系为例,分析物种丰富度和系统发育多样性的相关性。结果表明,蜈支洲岛植被可分为10种群系类型,22个样方的物种丰富度与系统发育多样性呈显著正相关,但系统发育多样性与物种丰富度对植物多样性保护优先顺序的指示有差异。森林群系的物种丰富度和系统发育多样性普遍高于灌木和草本群系,4个森林群系样方表现为系统发育结构发散,3个森林群系样方表现为系统发育结构聚集。灌木群系中露兜树(Pandanus tectorius)灌丛和草海桐(Scaevola sericea)灌丛系统发育多样性较高,草本群系中厚藤(Vitex trifolia)灌草丛系统发育多样性较高,灌木和草本群系大多倾向于系统发育结构聚集。距海或人类活动区较近的群系可能受环境过滤的作用,而远离海和人类活动区的群系主要受竞争排斥作用影响。因此,在岛屿植物多样性保护策略制定中应当综合考虑物种和系统发育维度,以及环境、演替阶段及用岛方式等因素的影响。     

Lake microbial communities are resilient after a whole-ecosystem disturbance

Disturbances act as powerful structuring forces on ecosystems. To ask whether environmental microbial communities have capacity to recover after a large disturbance event, we conducted a whole-ecosystem manipulation, during which we imposed an intense disturbance on freshwater microbial communities by artificially mixing a temperate lake during peak summer thermal stratification. We employed environmental sensors and water chemistry analyses to evaluate the physical and chemical responses of the lake, and bar-coded 16S ribosomal RNA gene pyrosequencing and automated ribosomal intergenic spacer analysis (ARISA) to assess the bacterial community responses. The artificial mixing increased mean lake temperature from 14 to 20 °C for seven weeks after mixing ended, and exposed the microorganisms to very different environmental conditions, including increased hypolimnion oxygen and increased epilimnion carbon dioxide concentrations. Though overall ecosystem conditions remained altered (with hypolimnion temperatures elevated from 6 to 20 °C), bacterial communities returned to their pre-manipulation state as some environmental conditions, such as oxygen concentration, recovered. Recovery to pre-disturbance community composition and diversity was observed within 7 (epilimnion) and 11 (hypolimnion) days after mixing. Our results suggest that some microbial communities have capacity to recover after a major disturbance.     

Distribution of Daphnia in a trade-off between food and temperature: individual habitat choice and time allocation

1. In a thermally stratified water column with a deep‐water algal maximum, Daphnia face a trade‐off between food (high fecundity) and temperature (fast development). Recent studies showed that Daphnia populations move up and down the entire water column to take advantage of both, but the proportion of time allocated by individuals to the epilimnion, metalimnion and hypolimnion with their specific food and temperature conditions is not yet known. 2. In a system of 1 m deep, vertical perspex tubes, I established three temperature gradients with 2, 5 and 10 °C differences between the surface (epilimnion) and the bottom layer (hypolimnion). Algae were added to the hypolimnion to simulate a deep‐water algal maximum. 3. The migration behaviour of individual neonate and egg‐bearing Daphnia hyalina × galeata was monitored in order to measure the proportions of time the individuals allocated to the different vertical habitats and to assess the frequency of their shifts between epilimnion and hypolimnion. 4. Neonates stayed continuously at the surface, taking advantage of the higher temperature, possibly because feeding was less important for them because of egg yolk reserves. In contrast, egg‐bearing females spent more time feeding in the hypolimnion when the temperature gradient was weak, but also migrated into the epilimnion to take advantage of the higher temperature. In the steepest temperature gradient, the egg‐bearing females either shifted between epilimnion and hypolimnion, or dwelled constantly in the metalimnion with intermediate conditions.     

The vertical distribution of some ciliated Protozoa in the plankton of a eutrophic pond during summer stratification

The vertical distribution of some ciliated Protozoa in the plankton of a pond in north-west England was investigated during August 1971. At this time, when the pond was stratified with an oxygen dificient hypolimnion, ciliates were counted at 10-cm depth intervals every 5 h over 25 h. The most common species (Loxodes magnus and L. striatus) were confined to the hypolimnion; there was no diurnal migration into the epilimnion. Earlier work had shown that Loxodes species require oxygen; it is therefore possible that these ciliates, which inhabited the oxygen dificient hypolimnion, migratedvertically, from time to time, to an oxygen supply at the boundary with the well-oxygenated epilimnion. To test this, Loxodes populations were confined in cellophane tubese both in the hypolimnion (at 3 m) and epilimnion (0.5 m) for 12 and 24 h (earlier trials had shown that the tubes were not markedly toxic). The ciliates died at both depths, and in a further experiment when Loxodes were confined at 3 m and 0.5 m and sampled at 5-h intervals up to 25 h it was found that they survived longer in the hypolimnion. It is suggested that ciliatees confined at 3 m died because they were unable to migrate vertically to an oxygen supply, while those at 0.5 m died because some other adverse factor was operating in the eiplimnion. Laboratory experiments showed that Loxodes died inn water in which phytoplankton photosynthesis took place and it is suggested that side effects of photosynthesis in the epilimnion (e.g. a rise in pH) caused the death of ciliates exposed at 0.5 m.     

Phylogenetic alpha and beta diversity in tropical tree assemblages along regional-scale environmental gradients in northwest South America

Aims Environmental gradients are drivers of species diversity; however, we know relatively little about the evolutionary processes underlying these relationships. A potentially powerful approach to studying diversity gradients is to quantify the phylogenetic structure within and between assemblages arrayed along broad spatial and environmental gradients. Here, we evaluate the phylogenetic structure of plant assemblages along an environmental gradient with the expectation that the habitat specialization of entire lineages is an important evolutionary pattern influencing the structure of tree communities along environmental gradients.Methods We evaluated the effect of several environmental variables on the phylogenetic structure of plant assemblages in 145 plots distributed in northwestern South America that cover a broad environmental gradient. The phylogenetic alpha diversity was quantified for each plot and the phylogenetic beta diversity between each pair of plots was also quantified. Both the alpha and beta diversity measures were then related to spatial and environmental gradients in the study system.Important findings We found that gradients in temperature and potential evapotranspiration have a strong relationship with the phylogenetic alpha diversity in our study system, with phylogenetic overdispersion in low temperatures and phylogenetic clustering at higher temperatures. Further, the phylogenetic beta diversity between two plots increases with an increasing difference in temperature, whereas annual precipitation was not a significant predictor of community phylogenetic turnover. We also found that the phylogenetic structure of the plots in our study system was related to the degree of seasonal flooding and seasonality in precipitation. In particular, more stressful environments such as dry forests and flooded forests showed phylogenetic clustering. Finally, in contrast with previous studies, we find that phylogenetic beta diversity was not strongly related to the spatial distance separating two forest plots, which may be the result of the importance of the three independent mountain ranges in our study system, which generate a high degree of environmental variation over very short distances. In conclusion, we found that environmental gradients are important drivers of both phylogenetic alpha and phylogenetic beta diversities in these forests over spatial distance.     

Morphological Characterization of Viruses in the Stratified Water Column of Alkaline, Hypersaline Mono Lake

Concentrations of viruses and prokaryotes in the alkaline, moderately hypersaline, seasonally stratified Mono Lake are among the highest reported for a natural aquatic environment. We used electron microscopy to test whether viral morphological characteristics differed among the epilimnion, metalimnion, and the anoxic hypolimnion of the lake and to determine how the properties of viruses in Mono Lake compare to other aquatic environments. Viral capsid size distributions were more similar in the metalimnion and hypolimnion of Mono Lake, while viral tail lengths were more similar in the epilimnion and metalimnion. The percentage of tailed viruses decreased with depth and the relative percentages of tailed phage families changed with depth. The presence of large (>125 nm capsid), untailed viruses in the metalimnion and hypolimnion suggests that eukaryotic viruses are produced in these suboxic and anoxic, hypersaline environments. Capsid diameters of viruses were larger on average in Mono Lake compared to other aquatic environments, and no lemon-shaped or filamentous viruses were found, in contrast to other high-salinity or high-altitude lakes and seas. Our data suggest that the physically and chemically distinct layers of Mono Lake harbor different viral assemblages, and that these assemblages are distinct from other aquatic environments that have been studied. Furthermore, we found that filtration of a sample through a 0.22-μm pore-size filter significantly altered the distribution of viral capsid diameters and tail lengths, resulting in a relative depletion of viruses having larger capsids and longer tails. This observation highlights the potential for bias in molecular surveys of viral diversity, which typically rely on filtration through 0.2- or 0.22-μm pore-size membrane filters to remove bacteria during sample preparation.     

Phylogenetic relatedness of co‐occurring waterbird communities: a test of Darwin's competition‐relatedness hypothesis

The competition–relatedness hypothesis of Darwin states that competition is greater among species that are phylogenetically closely‐related, and such species will tend to appear in separate communities (i.e. the species within communities will be phylogenetically overdispersed). Many studies have tested (and mainly refuted) this hypothesis for plant and bacterial communities. Results for the few studies with avian species are not conclusive. We tested Darwin's hypothesis for waterbirds using a set of open, artificial fish ponds in Doñana, south‐western Spain, that provide relatively homogeneous habitat where competition is likely to be intense. Monthly counts of 38 ponds (for 11 months, i.e. 418 censuses) recorded 76 bird species. Darwin's hypothesis predicted that species appearing in the same pond would be less related phylogenetically than expected if species occurred randomly across ponds and months according to the structure of the overall community across the entire pond complex. However, the waterbird community did not show a predominantly overdispersed pattern, suggesting that interspecific competition among phylogenetically related species was not the main force structuring communities. In contrast, the proportion of clustered communities was higher than expected throughout the annual cycle, indicating that related waterbirds tend to co‐occur on the same site, probably because they have similar microhabitat preferences. Clustering patterns were mainly driven by abundant and closely related duck species, and also by shorebirds. However, few individual pond communities remained significantly different from random after correction for multiple testing. Furthermore, the probability of co‐occurrence of a given species pair was negatively related to the phylogenetic distance between them. In conclusion, our study shows waterbird communities are mainly phylogenetically clustered or random, and do not support the competition‐relatedness hypothesis.     

沿海拔梯度变化的哀牢山亚热带森林群落系统发育结构

为揭示森林群落系统发育结构在海拔梯度上的变化及其驱动因素, 本研究以云南哀牢山西坡的亚热带森林群落为研究对象, 以APG III系统为基础框架, 结合DNA条形码序列信息解决末端分类单元亲缘关系的方法, 构建了哀牢山森林群落系统发育进化树, 采用净亲缘指数(net relatedness index, NRI)和最近亲缘指数(nearest taxon index, NTI), 探讨了不同植被类型的森林群落系统发育结构和沿海拔梯度的变化规律。结果表明, 从整体的海拔变化趋势上来看, 哀牢山森林群落系统发育结构随海拔上升由系统发育聚集(phylogenetic clustering)走向发散(phylogenetic overdispersion)或聚集程度降低。在低海拔地区, 群落表现为系统发育聚集, 表明生态位理论中的生境过滤作用在群落构建和生物多样性的维持中起着主导作用; 在中海拔地区, 出现了聚集与发散两种群落系统发育结构并存的现象, 推测可能是生境过滤和竞争排斥两种生态过程共同作用的结果; 在高海拔地区, 群落的系统发育结构因选择的指数不同而出现相反的结果, NRI表现出系统发育聚集, 而NTI却表现为随机或发散, 考虑到高海拔地区的环境胁迫可能促使植物发生趋同进化, 推测其群落构建的生态学过程需要更为综合的研究。本研究揭示群落系统发育结构沿海拔梯度确实存在显著的变化, 证明在生态群落的构建过程中非随机过程起到促进乃至关键作用。     

Changes in bacterioplankton community structure and activity with depth in a eutrophic lake as revealed by 5S rRNA analysis

The community structure of bacterioplankton was studied at different depths (0 to 25 m) of a temperate eutrophic lake (Lake Plusssee in northern Germany) by using comparative 5S rRNA analysis. The relative amounts of taxonomic groups were estimated from 5S rRNA bands separated by high-resolution electrophoresis. Comparison of partial 5S rRNA sequences enabled detection of changes in single taxa over space and during seasons. Overall, the bacterioplankton community was dominated by 3 to 14 abundant (>4% of the total 5S rRNA) taxa. In general, the number of 5S rRNA bands (i.e., the number of bacterial taxa) decreased with depth. In the fall, when thermal stratification and chemical stratification were much more pronounced than they were in the spring, the correlation between the depth layers and the community structure was more pronounced. Therefore, in the fall each layer had its own community structure; i.e., there were different community structures in the epilimnion, the metalimnion, and the hypolimnion. Only three 5S rRNA bands were detected in the hypolimnion during the fall, and one band accounted for about 70% of the total 5S rRNA. The sequences of individual 5S rRNA bands from the spring and fall were different for all size classes analyzed except two bands, one of which was identified as Comamonas acidivorans. In the overall analysis of the depth profiles, the diversity in the epilimnion contrasted with the reduced diversity of the bacterioplankton communities in the hypolimnion, and large differences occurred in the composition of the epilimnion at different seasons except for generalists like C. acidivorans.     

Environmental filtering is the main assembly rule of ground beetles in the forest and its edge but not in the adjacent grassland

In a fragmented landscape,transitional zones between neighboring habitats are common,and our understanding of community organizational forces across such habitats is important.Edge studies are numerous,but the majority of them utilize information on species richness and abundance.Abundance and taxonomic diversity,however,provide little information on the functioning and phylogeny of the co-existing species.Combining the evaluation of their functional and phylogenetic relationships,we aimed to assess whether ground beetle assemblages are deterministically or stochastically structured along grassland-forest gradients.Our results showed different community assembly rules on opposite sides of the forest edge.In the grassland,co-occurring species were functionally and phylogenetically not different from the random null model,indicating a random assembly process.Contrary to this,at the forest edge and the interior,co-occurring species showed functional and phylogenetic clustering,thus environmental filtering was the likely process structuring carabid assemblages.Community assembly in the grassland was considerably affected by asymmetrical species flows (spillover)across the forest edge:more forest species penetrated into the grassland than open-habitat and generalist species entered into the forest.This asymmetrical species flow underlines the importance of the filter function of forest edges.As unfavorable,human-induced changes to the structure,composition and characteristics of forest edges may alter their filter function,edges have to be specifically considered during conservation management.     

Fire modifies the phylogenetic structure of soil bacterial co‐occurrence networks

Fire alters ecosystems by changing the composition and community structure of soil microbes. The phylogenetic structure of a community provides clues about its main assembling mechanisms. While environmental filtering tends to reduce the community phylogenetic diversity by selecting for functionally (and hence phylogenetically) similar species, processes like competitive exclusion by limiting similarity tend to increase it by preventing the coexistence of functionally (and phylogenetically) similar species. We used co‐occurrence networks to detect co‐presence (bacteria that co‐occur) or exclusion (bacteria that do not co‐occur) links indicative of the ecological interactions structuring the community. We propose that inspecting the phylogenetic structure of co‐presence or exclusion links allows to detect the main processes simultaneously assembling the community. We monitored a soil bacterial community after an experimental fire and found that fire altered its composition, richness and phylogenetic diversity. Both co‐presence and exclusion links were more phylogenetically related than expected by chance. We interpret such a phylogenetic clustering in co‐presence links as a result of environmental filtering, while that in exclusion links reflects competitive exclusion by limiting similarity. This suggests that environmental filtering and limiting similarity operate simultaneously to assemble soil bacterial communities, widening the traditional view that only environmental filtering structures bacterial communities.     

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).