Canopy phylogenetic, chemical and spectral assembly in a lowland Amazonian forest

? Canopy chemistry and spectroscopy offer insight into community assembly and ecosystem processes in high-diversity tropical forests, but phylogenetic and environmental factors controlling chemical traits underpinning spectral signatures remain poorly understood. ? We measured 21 leaf chemical traits and spectroscopic signatures of 594 canopy individuals on high-fertility Inceptisols and low-fertility Ultisols in a lowland Amazonian forest. The spectranomics approach, which explicitly connects phylogenetic, chemical and spectral patterns in tropical canopies, provided the basis for analysis. ? Intracrown and intraspecific variation in chemical traits varied from 1.4 to 36.7% (median 9.3%), depending upon the chemical constituent. Principal components analysis showed that 14 orthogonal combinations were required to explain 95% of the variation among 21 traits, indicating the high dimensionality of canopy chemical signatures among taxa. Inceptisols and lianas were associated with high leaf nutrient concentrations and low concentrations of defense compounds. Independent of soils or plant habit, an average 70% (maximum 89%) of chemical trait variation was explained by taxonomy. At least 10 traits were quantitatively linked to remotely sensed signatures, which provided highly accurate species classification. ? The results suggest that taxa found on fertile soils carry chemical portfolios with a deep evolutionary history, whereas taxa found on low-fertility soils have undergone trait evolution at the species level. Spectranomics provides a new connection between remote sensing and community assembly theory in high-diversity tropical canopies.     

Environmental determinants of leaf litter ant community composition along an elevational gradient

Ant communities are extremely diverse and provide a wide variety of ecological functions in tropical forests. Here, we investigated the abiotic factors driving ant composition turnover across an elevational gradient at Mont Itoupé, French Guiana. Mont Itoupé is an isolated mountain whose top is covered by cloud forests, a biogeographical rarity that is likely to be threatened according to climate change scenarios in the region. We examined the influence of six soil, climatic, and LiDAR‐derived vegetation structural variables on leaf litter ant assembly (267 species) across nine 0.12‐ha plots disposed at three elevations (ca. 400, 600, and 800m asl). We tested (a) whether species cooccurring within a same plot or a same elevation were more similar in terms of taxonomic, functional, and phylogenetic composition, than species from different plots/elevations, and (b) which environmental variables significantly explained compositional turnover among plots. We found that the distribution of species and traits of ant communities along the elevational gradient was significantly explained by a turnover of environmental conditions, particularly in soil phosphorus and sand content, canopy height, and mean annual relative humidity of soil. Our results shed light on the role exerted by environmental filtering in shaping ant community assembly in tropical forests. Identifying the environmental determinants of ant species distribution along tropical elevational gradients could help predicting the future impacts of global warming on biodiversity organization in vulnerable environments such as cloud forests.     

A 7000‐year history of changing plant trait composition in an Amazonian landscape; the role of humans and climate

Tropical forests are shifting in species and trait composition, but the main underlying causes remain unclear because of the short temporal scales of most studies. Here, we develop a novel approach by linking functional trait data with 7000 years of forest dynamics from a fossil pollen record of Lake Sauce in the Peruvian Amazon. We evaluate how climate and human disturbances affect community trait composition. We found weak relationships between environmental conditions and traits at the taxon level, but strong effects for community‐mean traits. Overall, community‐mean traits were more responsive to human disturbances than to climate change; human‐induced erosion increased the dominance of dense‐wooded, non‐zoochorous species with compound leaves, and human‐induced fire increased the dominance of tall, zoochorous taxa with large seeds and simple leaves. This information can help to enhance our understanding of forest responses to past environmental changes, and improve predictions of future changes in tropical forest composition.     

Quantifying relationships between traits and explicitly measured gradients of stress and disturbance in early successional plant communities

Questions: How can one explicitly quantify, and separately measure, stress and disturbance gradients? How do these gradients affect functional composition in early successional plant communities and to what extent? Can we accurately predict trait composition from knowledge of these gradients? Location: Southern Quebec, Canada. Methods: Using eight environmental variables measured in 48 early successional plant communities, we estimated stress and disturbance gradients through structural equation modelling. We then measured 10 functional traits on the most abundant species of these 48 communities and calculated their community‐level mean and variance weighted by the relative abundance of each species. Finally, we related these community‐weighted means and variances to the estimated stress and disturbance gradients using general linear models or generalized additive models. Results: We obtained a well‐fitting measurement model of the stress and disturbance gradients existing in our sites. Of the 10 studied traits, only average plant reproductive height was strongly correlated with the stress (r²=0.464) and disturbance (r²=0.543) gradients. Leaf traits were not significantly related to either the stress or disturbance gradients. Conclusions: The well‐fitting measurement model of the stress and disturbance gradients, combined with the generally weak trait–environment linkages, suggests that community assembly in these early successional plant communities is driven primarily by stochastic processes linked to the history of arrival of propagules and not to trait‐based environmental filtering.     

Convergent elevation trends in canopy chemical traits of tropical forests

The functional biogeography of tropical forests is expressed in foliar chemicals that are key physiologically based predictors of plant adaptation to changing environmental conditions including climate. However, understanding the degree to which environmental filters sort the canopy chemical characteristics of forest canopies remains a challenge. Here, we report on the elevation and soil‐type dependence of forest canopy chemistry among 75 compositionally and environmentally distinct forests in nine regions, with a total of 7819 individual trees representing 3246 species collected, identified and assayed for foliar traits. We assessed whether there are consistent relationships between canopy chemical traits and both elevation and soil type, and evaluated the general role of phylogeny in mediating patterns of canopy traits within and across communities. Chemical trait variation and partitioning suggested a general model based on four interconnected findings. First, geographic variation at the soil‐Order level, expressing broad changes in fertility, underpins major shifts in foliar phosphorus (P) and calcium (Ca). Second, elevation‐dependent shifts in average community leaf dry mass per area (LMA), chlorophyll, and carbon allocation (including nonstructural carbohydrates) are most strongly correlated with changes in foliar Ca. Third, chemical diversity within communities is driven by differences between species rather than by plasticity within species. Finally, elevation‐ and soil‐dependent changes in N, LMA and leaf carbon allocation are mediated by canopy compositional turnover, whereas foliar P and Ca are driven more by changes in site conditions than by phylogeny. Our findings have broad implications for understanding the global ecology of humid tropical forests, and their functional responses to changing climate.     

Functional traits of individual trees reveal ecological constraints on community assembly in tropical rain forests

Niche differentiation and ecological filtering are primary ecological processes that shape community assembly, but their relative importance remains poorly understood. Analyses of the distributions of functional traits can provide insight into the community structure generated by these processes. We predicted the trait distributions expected under the ecological processes of niche differentiation and environmental filtering, then tested these predictions with a dataset of 4672 trees located in nine 1‐ha plots of tropical rain forest in French Guiana. Five traits related to leaf function (foliar N concentration, chlorophyll content, toughness, tissue density and specific leaf area), and three traits related to stem function (trunk sapwood density, branch sapwood density, and trunk bark thickness), as well as laminar surface area, were measured on every individual tree. There was far more evidence for environmental filtering than for niche differentiation in these forests. Furthermore, we contrasted results from species‐mean and individual‐level trait values. Analyses that took within‐species trait variation into account were far more sensitive indicators of niche differentiation and ecological filtering. Species‐mean analyses, by contrast, may underestimate the effects of ecological processes on community assembly. Environmental filtering appeared somewhat more intense on leaf traits than on stem traits, whereas niche differentiation affected neither strongly. By accounting for within‐species trait variation, we were able to more properly consider the ecological interactions among individual trees and between individual trees and their environment. In so doing, our results suggest that the ecological processes of niche differentiation and environmental filtering may be more pervasive than previously believed.     

Processes of community assembly in an environmentally heterogeneous,high biodiversity region

Despite decades of study, the relative importance of niche‐based versus neutral processes in community assembly remains largely ambiguous. Recent work suggests niche‐based processes are more easily detectable at coarser spatial scales, while neutrality dominates at finer scales. Analyses of functional traits with multi‐year multi‐site biodiversity inventories may provide deeper insights into assembly processes and the effects of spatial scale. We examined associations between community composition, species functional traits, and environmental conditions for plant communities in the Kouga‐Baviaanskloof region, an area within South Africa's Cape Floristic Region (CFR) containing high α and β diversity. This region contains strong climatic gradients and topographic heterogeneity, and is comprised of distinct vegetation classes with varying fire histories, making it an ideal location to assess the role of niche‐based environmental filtering on community composition by examining how traits vary with environment. We combined functional trait measurements for over 300 species with observations from vegetation surveys carried out in 1991/1992 and repeated in 2011/2012. We applied redundancy analysis, quantile regression, and null model tests to examine trends in species turnover and functional traits along environmental gradients in space and through time. Functional trait values were weakly associated with most spatial environmental gradients and only showed trends with respect to vegetation class and time since fire. However, survey plots showed greater compositional and functional stability through time than expected based on null models. Taken together, we found clear evidence for functional distinctions between vegetation classes, suggesting strong environmental filtering at this scale, most likely driven by fire dynamics. In contrast, there was little evidence of filtering effects along environmental gradients within vegetation classes, suggesting that assembly processes are largely neutral at this scale, likely the result of very high functional redundancy among species in the regional species pool.     

Examining variation in the leaf mass per area of dominant species across two contrasting tropical gradients in light of community assembly

Understanding variation in key functional traits across gradients in high diversity systems and the ecology of community changes along gradients in these systems is crucial in light of conservation and climate change. We examined inter‐ and intraspecific variation in leaf mass per area (LMA) of sun and shade leaves along a 3330‐m elevation gradient in Peru, and in sun leaves across a forest–savanna vegetation gradient in Brazil. We also compared LMA variance ratios (T‐statistics metrics) to null models to explore internal (i.e., abiotic) and environmental filtering on community structure along the gradients. Community‐weighted LMA increased with decreasing forest cover in Brazil, likely due to increased light availability and water stress, and increased with elevation in Peru, consistent with the leaf economic spectrum strategy expected in colder, less productive environments. A very high species turnover was observed along both environmental gradients, and consequently, the first source of variation in LMA was species turnover. Variation in LMA at the genus or family levels was greater in Peru than in Brazil. Using dominant trees to examine possible filters on community assembly, we found that in Brazil, internal filtering was strongest in the forest, while environmental filtering was observed in the dry savanna. In Peru, internal filtering was observed along 80% of the gradient, perhaps due to variation in taxa or interspecific competition. Environmental filtering was observed at cloud zone edges and in lowlands, possibly due to water and nutrient availability, respectively. These results related to variation in LMA indicate that biodiversity in species rich tropical assemblages may be structured by differential niche‐based processes. In the future, specific mechanisms generating these patterns of variation in leaf functional traits across tropical environmental gradients should be explored.     

Community structure of canopy arthropods associated to Quercus crassifolia×Quercus crassipes complex

Quercus crassifolia and Q. crassipes are dominant species in temperate forests of central Mexico and hybridize between each other when they occur in sympatry. Oak canopies contain a considerable portion of arthropod diversity and the hybrid zones can provide new habitats to epiphyte fauna. We tested if the establishment of hybrids in contact zones with their parental hosts increases the species diversity of canopy arthropods assuming that hybrid trees constitute new genotypes of potential new habitats to small organisms. We examined the effect of hybridization on some community structure parameters (diversity, composition, similarity and density of arthropod fauna) of canopy arthropods compared to their parental species in a hybrid zone located in central Mexico. We employed 17 leaf morphological traits and six diagnostic RAPD primers to identify parental and hybrid plants. The RAPDs marker showed unidirectional introgression towards Q. crassifolia, and were detected hybrid (F₁), backcrosses and introgression individual trees. In total, 30 oak canopies were fogged during rainy and dry season. We recognized 532 taxa of arthropods belonging to 22 orders associated with tree canopies. The taxonomic status of host‐trees may be an important factor in the arthropod community structure and that seasonality (dry and rainy) is not a factor that could modify their organization. Trees of Q. crassipes registered the highest densities of arthropod fauna followed by hybrid hosts (F₁); trees originated by backcrosses towards Q. crassifolia registered a significant less arthropod density than F₁ hybrids; and trees of Q. crassifolia had the lowest density. Hybrid plants and Q. crassipes individuals had higher diversity (H′) of arthropods than Q. crassifolia plants. Hybrid plants had also more rare species in both seasons in comparison with parental species. This study suggests that hybrid oaks act as a center of biodiversity by accumulating arthropods of both parental and different species including a considerable number of rare species.     

Can spatial variation in epiphyte diversity and community structure be predicted from sampling vascular epiphytes alone?

Aim Non‐vascular epiphytes have been largely ignored in studies examining the biotic and abiotic determinants of spatial variation in epiphyte diversity. Our aim was to test whether the spatial patterning of species richness, biomass and community composition across geographic regions, among trees within regions, and among branches within trees is consistent between the vascular and non‐vascular components of the temperate rain forest flora. Location Coastal lowland podocarp‐broadleaved forests on the west coast of the South Island of New Zealand. Methods We collected single samples (30 × 25 cm) from 96 epiphyte assemblages located on the inner branches of 40 northern rata (Metrosideros robusta) trees. For each sample, branch characteristics such as branch height, branch diameter, branch angle, branch aspect, and minimum and maximum epiphyte mat depth were recorded. The biomass for each individual epiphyte species was determined. Results Northern rata was host to a total of 157 species, comprising 32 vascular and 125 non‐vascular species, with liverworts representing 41% of all species. Within epiphyte mats, the average total organic biomass of 3.5 kg m^?2 of branch surface area consisted largely of non‐living biomass and roots. Vascular and non‐vascular epiphytes showed strikingly different spatial patterns in species richness, biomass and composition between sites, among trees within sites, and among branches within trees, which could not be explained by the branch structural characteristics we measured. The two plant groups had no significant association in community composition (r = 0.04, P = 0.08). However, the species richness of vascular plant seedlings was strongly linked to the presence/absence of lichens. Main conclusions Non‐vascular plants contributed substantially to the high species richness and biomass recorded in this study, which was comparable to that of some tropical rain forests. High variability in community composition among epiphyte mats, and very low correlation with any of the environmental factors measured possibly indicate high levels of stochasticity in seed or spore colonization, establishment success or community assembly among branches in these canopy communities. Although we found some evidence that vascular plant seedling establishment was linked to the presence of lichens and the biomass of non‐living components in the epiphyte mats, there was no correlation in the spatial patterning or determinants of species richness between non‐vascular and vascular plants. Consequently, variation in total epiphyte biodiversity could not be predicted from the measurement of vascular plant diversity alone, which highlights the crucial importance of sampling non‐vascular plants when undertaking epiphyte community studies.     

Response of floodplain understorey species to environmental gradients and tree invasion: a functional trait perspective

Plants are connected to habitats by functional traits which are filtered by environmental gradients. Since tree species composition in the forest canopy can influence ecosystem processes by changing resource availability, litter accumulation, and soil nutrient content, we hypothesised that non-native invasive trees can establish new environmental filters on the understorey communities. In the hardwood floodplain forests in Northern Italy, the invasive trees Robinia pseudoacacia L. and Prunus serotina Ehrh. are the dominant canopy species. We used trait data assembled from databases and iterative RLQ analysis to identify a parsimonious set of functional traits responding to environmental variables (soil, light availability, disturbance, and stand structure) and the dominant native and invasive canopy species. Then, RLQ and fourth-corner analysis was conducted to investigate the joint structure between macro-environmental variables and species traits and functional groups were identified. The trait composition of the herb-layer was significantly related to the main environmental gradients and the presence of the invaders in the canopy showed significant relationships with several traits. In particular, the presence of P. serotina may mitigate or even erase the effect of disturbances, maintaining a stable forest microclimate and thus favouring ‘true’ forest species, while R. pseudoacacia may slow down forest succession and regeneration by establishing new stable associations with a graminoid-dominated understorey. The impact of the two invasive trees on herb layer composition appears to differ, indicating that different management and control strategies may be needed.     

Underdispersion and overdispersion of traits in terrestrial snail communities on islands

Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under‐ and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM‐RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small‐scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.     

Among‐species overlap in rodent body size distributions predicts species richness along a temperature gradient

Temperature is widely regarded as a major driver of species richness, but the mechanisms are debated. Niche theory suggests temperature may affect richness by filtering traits and species in colder habitats while promoting specialization in warmer ones. However, tests of this theory are rare because niche dimensions are challenging to quantify along broad thermal gradients. Here, we use individual‐level trait data from a long‐term monitoring network spanning a large geographic extent to test niche‐based theory of community assembly in small mammals. We examined variation in body size among 23 communities of North American rodents sampled across the National Ecological Observatory Network (NEON), ranging from northern hardwood forests to subtropical deserts. We quantified body size similarity among species using a metric of overlap that accounts for individual variation, and fit a structural equation model to disentangle the relationships between temperature, productivity, body size overlap, and species richness. We document a latitudinal gradient of declining similarity in body size among species towards the tropics and overall increase in the dimensions of community‐wide trait space in warmer habitats. Neither environmental temperature nor net primary productivity directly affect rodent species richness. Instead, temperature determines the community‐wide niche space that species can occupy, which in turn alters richness. We suggest a latitudinal gradient of trait space expansion towards the tropics may be widespread and underlie gradients in species diversity.     

Expanding the understanding of local community assembly in adaptive radiations

Communities are thought to be assembled by two types of filters: by the environment relating to the fundamental niche and by biotic interactions relating to the realized niche. Both filters include parameters related to functional traits and their variation along environmental gradients. Here, we infer the general importance of environmental filtering of a functional trait determining local community assembly within insular adaptive radiations on the example of Caribbean Anolis lizards. We constructed maps for the probability of presence of Anolis ecomorphs (ecology‐morphology‐behavior specialists) on the Greater Antilles and overlaid these to estimate ecomorph community completeness (ECC) over the landscape. We then tested for differences in environmental parameter spaces among islands for real and cross‐fitted ECC values to see whether the underlying assembly filters are deterministic (i.e., similar among islands). We then compared information‐theoretic models of climatic and landscape parameters among Greater Antillean islands and inferred whether body mass as functional trait determines ECC. We found areas with high ECC to be strongly correlated with environmental filters, partly related to elevation. The environmental parameters influencing high ECC differed among islands. With the exception of the Jamaican twig ecomorph (which we suspect to be misclassified), smaller ecomorphs were more restricted to higher elevations than larger ones which might reflect filtering on the basis of differential physiological restrictions of ecomorphs. Our results in Anolis show that local community assembly within adaptive island radiations of animals can be determined by environmental filtering of functional traits, independently from species composition and realized environmental niche space.     

Cryptogamic epiphytes in primary and recovering upper montane oak forests of Costa Rica – species richness,community composition and ecology

Species richness, community composition and ecology of cryptogamic epiphytes (bryophytes, macrolichens) were studied in upper montane primary, early secondary and late secondary oak forests of the Cordillera de Talamanca, Costa Rica. Canopy trees of Quercus copeyensis were sampled with the aim of getting insight in patterns and processes of epiphyte succession and recovery of diversity in secondary forest following forest clearing. Species richness of cryptogamic epiphytes in secondary and primary forests were nearly the same, showing that primary forests are not necessarily more diverse than secondary forests. High species richness of secondary forests was presumed due to the closed canopy, resulting in permanently high atmospheric humidity in these forests. Similarity in species composition of secondary and primary forests increases with forest age, but after 40 years of succession one third (46 species) of primary forest species had not re-established in the secondary forest. Community composition in primary and secondary forests differed markedly and indicates that a long time is needed for the re-establishment of microhabitats and re-invasion of species and communities adapted to differentiated niches. Genera and species exclusive to primary forests are relevant as indicator taxa and conservation targets. Forty percent (68 species) of all species recorded are restricted to secondary forests, indicating the important contribution of secondary forest diversity to total species richness of the oak forests of Costa Rica.     

Greater variation in boreal plant community composition and community-level traits at local- than regional-scale

Questions

Rapid climate change in northern latitudes is expected to influence plant functional traits of the whole community (community-level traits) through species compositional changes and/or trait plasticity, limiting our ability to anticipate climate warming impacts on northern plant communities. We explored differences in plant community composition and community-level traits within and among four boreal peatland sites and determined whether intra- or interspecific variation drives community-level traits.

Location

Boreal biome of western North America.

Methods

We collected plant community composition and functional trait data along dominant topoedaphic and/or hydrologic gradients at four peatland sites spanning the latitudinal extent of the boreal biome of western North America. We characterized variability in community composition and community-level traits of understorey vascular and moss species both within (local-scale) and among sites (regional-scale).

Results

Against expectations, community-level traits of vascular plant and moss species were generally consistent among sites. Furthermore, interspecific variation was more important in explaining community-level trait variation than intraspecific variation. Within-site variation in both community-level traits and community composition was greater than among-site variation, suggesting that local environmental gradients (canopy density, organic layer thickness, etc.) may be more influential in determining plant community processes than regional-scale gradients.

Conclusions

Given the importance of interspecific variation to within-site shifts in community-level traits and greater variation of community composition within than among sites, we conclude that climate-induced shifts in understorey community composition may not have a strong influence on community-level traits in boreal peatlands unless local-scale environmental gradients are substantially altered.     

Diversity patterns of vascular epiphytes along an elevational gradient in the Andes

Aim To document the elevational pattern of epiphyte species richness at the local scale in the tropical Andes with a consistent methodology. Location The northern Bolivian Andes at 350–4000 m above sea level. Methods We surveyed epiphytic vascular plant assemblages in humid forests in (a) single trees located in (b) 90 subplots of 400 m² each located in (c) 14 plots of 1 ha each. The plots were separated by 100–800 m along the elevational gradient. Results We recorded about 800 epiphyte species in total, with up to 83 species found on a single tree. Species richness peaked at c. 1500 m and declined by c. 65% to 350 m and by c. 99% to 4000 m, while forests on mountain ridges had richness values lowered by c. 30% relative to slope forests at the same elevations. The hump‐shaped richness pattern differed from a null‐model of random species distribution within a bounded domain (the mid‐domain effect) as well as from the pattern of mean annual precipitation by a shift of the diversity peak to lower elevations and by a more pronounced decline of species richness at higher elevations. With the exception of Araceae, which declined almost monotonically, all epiphyte taxa showed hump‐shaped curves, albeit with slightly differing shapes. Orchids and pteridophytes were the most species‐rich epiphytic taxa, but their relative contributions shifted with elevation from a predominance of orchids at low elevations to purely fern‐dominated epiphyte assemblages at 4000 m. Within the pteridophytes, the polygrammoid clade was conspicuously overrepresented in dry or cold environments. Orchids, various small groups (Cyclanthaceae, Ericaceae, Melastomataceae, etc.), and Bromeliaceae (below 1000 m) were mostly restricted to the forest canopy, while Araceae and Pteridophyta were well represented in the forest understorey. Main conclusions Our study confirms the hump‐shaped elevational pattern of vascular epiphyte richness, but the causes of this are still poorly understood. We hypothesize that the decline of richness at high elevations is a result of low temperatures, but the mechanism involved is unknown. The taxon‐specific patterns suggest that some taxa have a phylogenetically determined propensity for survival under extreme conditions (low temperatures, low humidity, and low light levels in the forest interior). The three spatial sampling scales show some different patterns, highlighting the influence of the sampling methodology.     

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.     

Phylogenetic community structure and phylogenetic turnover across space and edaphic gradients in western Amazonian tree communities

Ecological and evolutionary processes influence community assembly at both local and regional scales. Adding a phylogenetic dimension to studies of species turnover allows tests of the extent to which environmental gradients, geographic distance and the historical biogeography of lineages have influenced speciation and dispersal of species throughout a region. We compare measures of beta diversity, phylogenetic community structure and phylobetadiversity (phylogenetic distance among communities) in 34 plots of Amazonian trees across white‐sand and clay terra firme forests in a 60 000 square kilometer area in Loreto, Peru. Dominant taxa in white‐sand forests were phylogenetically clustered, consistent with environmental filtering of conserved traits. Phylobetadiversity measures found significant phylogenetic clustering between terra firme communities separated by geographic distances of <200–300 km, consistent within recent local speciation at the watershed scale in the Miocene‐aged clay‐soil forests near the foothills of the Andes. Although both distance and habitat type yielded statistically significant effects on both species and phylogenetic turnover, the patterns we observed were more consistent with an effect of habitat specialization than dispersal limitation. Our results suggest a role for both broad‐scale biogeographic and evolutionary processes, as well as habitat specialization, influencing community structure in Amazonian forests.     

Plant Trait Variation along an Altitudinal Gradient in Mediterranean High Mountain Grasslands: Controlling the Species Turnover Effect

Assessing changes in plant functional traits along gradients is useful for understanding the assembly of communities and their response to global and local environmental drivers. However, these changes may reflect the effects of species composition (i.e. composition turnover), species abundance (i.e. species interaction), and intra-specific trait variability (i.e. species plasticity). In order to determine the relevance of the latter, trait variation can be assessed under minimal effects of composition turnover. Nine sampling sites were established along an altitudinal gradient in a Mediterranean high mountain grassland community with low composition turnover (Madrid, Spain; 1940 m–2419 m). Nine functional traits were also measured for ten individuals of around ten plant species at each site, for a total of eleven species across all sites. The relative importance of different sources of variability (within/between site and intra-/inter-specific functional diversity) and trait variation at species and community level along the considered gradients were explored. We found a weak individual species response to altitude and other environmental variables although in some cases, individuals were smaller and leaves were thicker at higher elevations. This lack of species response was most likely due to greater within- than between-site species variation. At the community level, inter-specific functional diversity was generally greater than the intra-specific component except for traits linked to leaf element content (leaf carbon content, leaf nitrogen content, δ¹³C and δ¹⁵N). Inter-specific functional diversity decreased with lower altitude for four leaf traits (specific leaf area, leaf dry matter content, δ¹³C and δ¹⁵N), suggesting trait convergence between species at lower elevations, where water shortage may have a stronger environmental filtering effect than colder temperatures at higher altitudes. Our results suggest that, within a vegetation type encompassing various environmental gradients, both, changes in species abundance and intra-specific trait variability adjust for the community functional response to environmental changes.