Patterns in a species‐rich tropical understory plant community

Understory plants are an important component of the high plant species diversity characteristic of neotropical rain forests. Herbs, shrubs, understory trees, and saplings of canopy trees occupy a broadly uniform environment of abundant rainfall, low light levels, and high humidity. We asked whether this community at the La Selva Biological Station in the Caribbean lowlands of Costa Rica was structured by environmental filters such as soil origin, topographic position, and understory light availability. We used nested quadrats to assess effects of soil origin (recent alluvium, weathered alluvium, residual volcanic soil) and topographic position (ridges, mid‐slopes and flats) on species composition, density, and diversity and measured six edaphic and understory light parameters. Non‐metric multidimensional scaling ordinations were based on frequency of occurrence in 20 quadrats for 272 species in the shrub size class and 136 species in the small‐tree size class for 17 sites. Three axes were correlated with composite environmental variables produced by principal component analysis representing slope, extractable phosphorus, and light. NMS site positions also reflected soil origin, topographic position, and geographic location. The analyses illustrated a complex community structured by species responses to environmental filters at multiple, interdigitated spatial scales. We suggest that light availability affected by canopy dynamics and dispersal limitation provides additional sources of variation in species distributions, which interact with edaphic patterns in complex ways. Abstract in Spanish is available with online material.     

Determinants of Tree Assemblage Composition at the Mesoscale within a Subtropical Eucalypt Forest

A variety of environmental processes, including topography, edaphic and disturbance factors can influence vegetation composition. The relative influence of these patterns has been known to vary with scale, however, few studies have focused on environmental drivers of composition at the mesoscale. This study examined the relative importance of topography, catchment flow and soil in influencing tree assemblages in Karawatha Forest Park; a South-East Queensland subtropical eucalypt forest embedded in an urban matrix that is part of the Terrestrial Ecosystem Research Network South-East Queensland Peri-urban SuperSite. Thirty-three LTER plots were surveyed at the mesoscale (909 ha), where all woody stems ≥1.3 m high rooted within plots were sampled. Vegetation was divided into three cohorts: small (≥1–10 cm DBH), intermediate (≥10–30 cm DBH), and large (≥30 cm DBH). Plot slope, aspect, elevation, catchment area and location and soil chemistry and structure were also measured. Ordinations and smooth surface modelling were used to determine drivers of vegetation assemblage in each cohort. Vegetation composition was highly variable among plots at the mesoscale (plots systematically placed at 500 m intervals). Elevation was strongly related to woody vegetation composition across all cohorts (R²: 0.69–0.75). Other topographic variables that explained a substantial amount of variation in composition were catchment area (R²: 0.43–0.45) and slope (R²: 0.23–0.61). Soil chemistry (R²: 0.09–0.75) was also associated with woody vegetation composition. While species composition differed substantially between cohorts, the environmental variables explaining composition did not. These results demonstrate the overriding importance of elevation and other topographic features in discriminating tree assemblage patterns irrespective of tree size. The importance of soil characteristics to tree assemblages was also influenced by topography, where ridge top sites were typically drier and had lower soil nutrient levels than riparian areas.     

Habitat Associations and Community Structure of Dipterocarps in Response to Environment and Soil Conditions in Brunei Darussalam,Northwest Borneo

Plant habitat associations are well documented in Bornean lowland tropical forests, but few studies contrast the prevalence of associations across sites. We examined habitat associations and community composition of Dipterocarpaceae trees in two contrasting Bornean lowland mixed dipterocarp forests separated by approximately 100 km: Andulau (uniform topography, lower altitudinal range, sandy soils) and Belalong (highly dissected topography, higher altitudinal range, clay‐rich soils). Dipterocarpaceae trees ≥ 1 cm diameter at breast height (dbh) were censused in 20‐m wide belt transects established along topographic gradients at each site. Dipterocarp density, evenness, species richness, and diversity were significantly higher at Andulau than Belalong. Significant site associations (with either Andulau or Belalong) were detected for 19 (52%) of the 37 dipterocarp species tested. Dipterocarpaceae community composition at Belalong correlated with soil nutrient concentrations as well as measures of vegetation and topographic structure, but community composition at Andulau correlated with fewer habitat variables. Within each site, dipterocarp density, species richness, and diversity were consistently higher on ridges than in slopes and valleys. Significant within‐site associations to topographic habitats were less common at Andulau (10% of species tested) than at Belalong (15%). We conclude that edaphic and other environmental factors influence dipterocarp community composition at a local scale, and are more important drivers of community structure in the more variable environment at Belalong. Species richness and diversity of dipterocarps on small plots, however, were higher at Andulau, suggesting that factors other than environmental heterogeneity contribute to contrasts in dipterocarp tree species richness at small scales.     

Environmental drivers of tree community turnover in western Amazonian forests

A more comprehensive understanding of the factors governing tropical tree community turnover at different spatial scales is needed to support land‐management and biodiversity conservation. We used new forest inventory data from 263 permanent plots in the Carnegie Biodiversity‐Biomass Forest Plot Network spanning the eastern Andes to the western Amazonian lowlands of Peru to examine environmental factors driving genus‐level canopy tree compositional variation at regional and landscape scales. Across the full plot network, constrained ordination analysis indicated that all environmental variables together explained 23.8% of the variation in community composition, while soil, topographic, and climatic variables each explained 15.2, 10.9, and 17.0%, respectively. A satellite‐derived metric of cloudiness was the single strongest predictor of community turnover, and constrained ordination revealed a primary gradient of environmentally‐driven community turnover spanning from cloudy, high elevation sites to warm, wet, lowland sites. For three focal landscapes within the region, local environmental variation explained 13.4–30.8% of compositional variation. Community turnover at the landscape scale was strongly driven by topo‐edaphic factors in the two lowland landscapes examined and strongly driven by potential insolation and topography in the montane landscape. At the regional scale, we found that the portion of compositional variation that was uniquely explained by spatial variation was relatively small (2.7%), and was effectively zero within the three focal landscapes. Overall, our results show strong canopy tree compositional turnover in response to environmental gradients at both regional and landscape scales, though the most important environmental drivers differed between scales and among landscapes. Our results also highlight the usefulness of key satellite‐derived environmental covariates that should be considered when conducting biodiversity analyses in tropical forests.     

Tropical metacommunities along elevational gradients: effects of forest type and other environmental factors

Elevational gradients provide a natural experiment for assessing the extent to which the structure of animal metacommunities is molded by biotic and abiotic characteristics that change gradually, or is molded by aspects of plant community composition and physiognomy that change in a more discrete fashion. We used a metacommunity framework to integrate species‐specific responses to environmental gradients as an approach to detect emergent patterns at the mesoscale in the Luquillo Mountains of Puerto Rico. Elements of metacommunity structure (coherence, species turnover and range boundary clumping) formed the basis for distinguishing among random, checkerboard, Gleasonian, Clementsian, evenly spaced and nested patterns. Paired elevational transects (300–1000 m a.s.l.) were sampled at 50 m intervals to decouple underlying environmental mechanisms: a mixed forest transect reflected changes in abiotic and biotic conditions, including forest type (i.e. tabonuco, palo colorado and elfin forests), whereas another transect reflected changes in environmental conditions but not forest type, as its constituent plots were located within palm forest. Based on distributional data (presence versus absence of species), the mixed forest transect exhibited Clementsian structure, whereas the palm forest transect exhibited quasi‐Gleasonian structure. In contrast, the distribution of modes in species abundance was random with respect to the latent environmental gradient in the mixed forest transect and clumped with respect to the latent environmental gradient in the palm forest transect. Such contrasts suggest that the environmental factors affecting abundance differed in form or type from those affecting distributional boundaries. Variation among elevational strata with respect to the first axis of correspondence from reciprocal averaging was highly correlated with elevation along each transect, even though axis scores were not correlated between mixed forest and palm forest transects. This suggests that the identity of the environmental characteristics, or the form of response by the fauna to those characteristics, differed between the two elevational transects. Despite the proximity of the transects, the patchy configuration of palm forest, and the pervasive distribution of the dominant palm species, the relative importance of abiotic variables and habitat in structuring gastropod metacommunities differed between transects, which is remarkable and attests to the sensitivity of metacommunity structure to environmental variation.     

Community Functional Responses to Soil and Climate at Multiple Spatial Scales: When Does Intraspecific Variation Matter?

Despite increasing evidence of the importance of intraspecific trait variation in plant communities, its role in community trait responses to environmental variation, particularly along broad-scale climatic gradients, is poorly understood. We analyzed functional trait variation among early-successional herbaceous plant communities (old fields) across a 1200-km latitudinal extent in eastern North America, focusing on four traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We determined the contributions of species turnover and intraspecific variation to between-site functional dissimilarity at multiple spatial scales and community trait responses to edaphic and climatic factors. Among-site variation in community mean trait values and community trait responses to the environment were generated by a combination of species turnover and intraspecific variation, with species turnover making a greater contribution for all traits. The relative importance of intraspecific variation decreased with increasing geographic and environmental distance between sites for SLA and leaf area. Intraspecific variation was most important for responses of vegetative height and responses to edaphic compared to climatic factors. Individual species displayed strong trait responses to environmental factors in many cases, but these responses were highly variable among species and did not usually scale up to the community level. These findings provide new insights into the role of intraspecific trait variation in plant communities and the factors controlling its relative importance. The contribution of intraspecific variation to community trait responses was greatest at fine spatial scales and along edaphic gradients, while species turnover dominated at broad spatial scales and along climatic gradients.     

Palm distribution patterns in Amazonian rainforests: What is the role of topographic variation?

Question: Are soil properties and topographic variation related with palm (Arecaceae) species composition and distribution patterns? If so, are species distribution patterns consistent across sites? Location: 100–200 m a.s.l, non‐inundated Amazonian rainforest, NE Peru. Methods: One 0.65‐ha line transect divided into 5 m by 5 m subunits was inventoried for all palm individuals at each site. Soil samples were collected, and topography was measured. Results: A total of 56 palm taxa were recorded. Floristic similarity among transects clearly corresponded with similarity in soil cation content when species abundances were taken into account, but less so when only presence‐absence data were used. Taxon‐wise distribution analyses were done for the 37 most abundant palm taxa. Quite a few of these taxa proved not to be randomly distributed along the transects, but instead were clearly more abundant in some topographic positions than in others. However, the consistency of the distribution patterns across study sites proved to be rather low, and many of the palm taxa showed different distribution patterns at different sites. Conclusions: The ambiguity in distribution patterns across study sites may partly be due to the complexity of topography as a measure of ecological conditions, and the probability that it is related to the variation in different environmental variables at different sites.     

A model of geographical, environmental and regional variation in vegetation composition of pyrogenic grasslands of Florida

Aim To develop a landscape‐level model that partitions variance in plant community composition among local environmental, regional environmental, and purely spatial predictive variables for pyrogenic grasslands (prairies, savannas and woodlands) throughout northern and central Florida. Location North and central Florida, USA. Methods We measured plant species composition and cover in 271 plots throughout the study region. A variation‐partitioning model was used to quantify components of variation in species composition associated with the main and interaction effects of soil and topographic variables, climate variables and spatial coordinates. Partial correlations of environmental variables with community variation were identified using direct gradient analysis (redundancy analysis and partial redundancy analysis) and Monte Carlo tests of significance. Results Community composition was most strongly related to edaphic variables at local scales in association with topographic gradients, although geographically structured edaphic, climatic and pure spatial effects were also evident. Edaphic variables explained the largest portion of total variation explained (TVE) as a main effect (48%) compared with the main effects of climate (9%) and pure spatial factors (9%). The remaining TVE was explained by the interaction effect of climate and spatial factors (13%) and the three‐way interaction (22%). Correlation analyses revealed that the primary compositional gradient was related to soil fertility and topographic position corresponding to soil moisture. A second gradient represented distinct geographical separation between the Florida panhandle and peninsular regions, concurrent with differences in soil characteristics. Gradients in composition corresponded to species richness, which was lower in the Florida peninsula. Main conclusions Environmental variables have the strongest influence on the species composition of Florida pyrogenic grasslands at both local and regional scales. However, the limited distributions of many plant taxa suggest historical constraints on species distributions from one physiographical region to the other (Florida panhandle and peninsula), although this pattern is partially confounded by regionally spatially structured environmental variables. Our model provides insight into the relative importance of local‐ and regional‐scale environmental effects as well as possible historical constraints on floristic variation in pine‐dominated pyrogenic grasslands of the south‐eastern USA.     

Soil‐based habitat partitioning in understorey palms in lower montane tropical forests

Aim Dispersal assembly and niche assembly are two competing theories proposed to explain the maintenance of species diversity in tropical forests. Dispersal theory emphasizes the role of chance colonization events and distance‐limited seed dispersal in explaining species abundance and distribution, whereas niche theory emphasizes differences among species in requirements for potentially limiting resources. Species distribution patterns in tropical forests often correlate with geology and topography, but tests of the relative importance of dispersal and niche partitioning have been hampered by an inadequate characterization of resource availability. The aim of this study was to explore how soil chemical and physical properties, climate, and geographic distance affect understorey palm communities in lower montane forests. Location Fortuna Forest Reserve, Chiriqui Province, and Palo Seco Forest Reserve, Bocas del Toro Province, in western Panama. Methods Understorey palms and soil nutrient concentrations were surveyed within 10 sites on different soil types across a 13‐km transect. Variation in palm community composition was examined in relation to spatial and environmental variables. Results The 25 understorey palm species recorded in the study were non‐randomly distributed among forests differing in soil nutrient availability. In support of dispersal theory, floristic similarity decreased predictably with increasing geographic distance. However, environmental and soil variables were also correlated with geographic distance. Floristic similarity was also highly associated with a subset of environmental variables. Variation in palm community similarity was most strongly correlated with inorganic nitrogen availability and cation concentration. A subset of soil variables had a stronger relationship with floristic similarity when geographic distance was controlled for than did geographic distance when differences in soils were controlled for. Main conclusions Both dispersal and niche processes affect palm species distribution patterns. Although spatially limited dispersal may influence species distribution patterns, soil‐based habitat associations, particularly with respect to soil nitrogen, cation availability and aluminium concentrations, remain important factors influencing palm community composition at the mesoscale level in this tropical montane forest.     

Contrasting altitudinal variation of alpine plant communities along the Swedish mountains

Changes in abiotic factors along altitudinal and latitudinal gradients cause powerful environmental gradients. The topography of alpine areas generates environmental gradients over short distances, and alpine areas are expected to experience greater temperature increase compared to the global average. In this study, we investigate alpha, beta, and gamma diversity, as well as community structure, of vascular plant communities along altitudinal gradients at three latitudes in the Swedish mountains. Species richness and evenness decreased with altitude, but the patterns within the altitudinal gradient varied between sites, including a sudden decrease at high altitude, a monotonic decrease, and a unimodal pattern. However, we did not observe a decline in beta diversity with altitude at all sites, and plant communities at all sites were spatially nested according to some other factors than altitude, such as the availability of water or microtopographic position. Moreover, the observed diversity patterns did not follow the latitudinal gradient. We observed a spatial modularity according to altitude, which was consistent across sites. Our results suggest strong influences of site‐specific factors on plant community composition and that such factors partly may override effects from altitudinal and latitudinal environmental variation. Spatial variation of the observed vascular plant communities appears to have been caused by a combination of processes at multiple spatial scales.     

Consistent Richness-Biomass Relationship across Environmental Gradients in a Marine Macroalgal-Dominated Subtidal Community on the Western Antarctic Peninsula

Biodiversity loss has spurred the biodiversity-ecosystem functioning research over a range of ecosystems. In Antarctica, however, the relationship of taxonomic and functional diversity with ecosystem properties (e.g., community biomass) has received less attention, despite the presence of sharp and dynamic environmental stress gradients that might modulate these properties. Here, we investigated whether the richness-biomass relationship in macrobenthic subtidal communities is still apparent after accounting for environmental stress gradients in Fildes Bay, King George Island, Antarctica. Measurements of biomass of mobile and sessile macrobenthic taxa were conducted in the austral summer 2013/4 across two environmental stress gradients: distance from nearest glaciers and subtidal depth (from 5 to 30 m). In general, community biomass increased with distance from glaciers and water depth. However, generalised additive models showed that distance from glaciers and depth accounted for negligible proportions of variation in the number of functional groups (i.e., functional richness) and community biomass when compared to taxonomic richness. Functional richness and community biomass were positive and saturating functions of taxonomic richness. Large endemic, canopy-forming brown algae of the order Desmarestiales dominated the community biomass across both gradients. Accordingly, differences in the composition of taxa accounted for a significant and large proportion (51%) of variation in community biomass in comparison with functional richness (10%). Our results suggest that the environmental factors here analysed may be less important than biodiversity in shaping mesoscale (several km) biomass patterns in this Antarctic system. We suggest that further manipulative, hypothesis-driven research should address the role of biodiversity and species’ functional traits in the responses of Antarctic subtidal communities to environmental variation.     

Local Hydrological Conditions Explain Floristic Composition in Lowland Amazonian Forests

Amazonian forests harbor a large variety of understory herbs adapted to areas with different hydrological conditions, ranging from well‐drained to seasonally flooded forests. The presence versus absence of flooding forms the extremes of a hydrological gradient, with various intermediate conditions, such as seasonal soil waterlogged areas, in between. We investigated the relationship between understory herbs and hydrological conditions in Central Amazonian forests using eighty‐eight 250 × 2 m plots distributed along a 600‐km transect. Hydrological conditions were determined regionally by precipitation and locally by topographic conditions based on drainage potential, flooding height and soil permeability (sand content). Soil cation concentration was used as a proxy for soil fertility. The floristic dissimilarities among plots were visualized by Nonmetric Multidimensional Scaling, and simple and multiple regressions were used to identify the best predictor of herb species composition. Local drainage potential was more important in determining herb species composition than soil fertility or precipitation at non‐flooded and flooded sites. Flooded sites comprised a very distinctive herb species composition even when the flood height was low (0.3 m). We conclude that hydrological conditions are the primary constraint of herb distribution within this flat regional landscape with moderate amounts of soil fertility variation (0.09–2.280 cmol(+)/kg). Hydrological models that consider local water conditions explained the largest part of herb species composition. Therefore, predictions of species distribution based on large‐scale climatic variables may underestimate the favorable area for understory herbs if the variation on local hydrological conditions is not considered.     

The influence of distance to perennial surface water on ant communities in Mopane woodlands,northern Botswana

Studies of biodiversity along environmental gradients provide information on how ecological communities change in response to biotic and abiotic factors. For instance, distance to water is associated with several factors that shape the structure and the functioning of ecosystems at a range of spatial scales. We investigated the influence of distance to a perennial water source on ant communities in a semi‐arid savanna in northern Botswana. Ant abundance, taxonomic richness, and both alpha and beta diversity were generally higher during the wet than the dry season. However, there were strong seasonal influences on the effects of distance to water, with more pronounced effects during the wet season. While both abundance and beta diversity declined with increasing distances to water during the wet season, there was a contrasting increase in alpha diversity. There was no major effect of distance to water on taxonomic richness during either season. Beta diversity was as high across as along gradients, and we found support for modular rather than nested community structures along gradients. Our study demonstrated that small‐scale gradients in distance to water can influence several aspects of ant communities in semi‐arid savannas. However, our results also point to strong effects of small‐scale environmental variation, for instance associated with vegetation characteristics, soil properties, and plant community structure that are not directly linked to water access.     

Mesoscale distribution of understorey plants in temperate forest (Kalø, Denmark): the importance of environment and dispersal

We studied the influence of environmental heterogeneity and dispersalprocesses on mesoscale distribution patterns of understorey plants in a 330ha ancient, managed temperate forest area. Similarity matrixregression analysis showed that overall species similarity was affected bysite(the two disjunct halves of the forest area), spatial autocorrelation at the100m scale, edaphic conditions, stand type and stand structure, andthe occurrence of open areas, but not by stand age or topography. Usingautologistic regression we analysed the influence of clumping, site, edaphicconditions, stand composition, open areas, and stand structure on theindividualdistribution patterns of the 60 most abundant species. Only five species haddistributions that were not significantly related to any of the explanatoryvariables. A large number of species showed significant, and often differing,relationships to at least one of the environmental parameters. At least 20species exhibited clumping at the 100 m scale that was independentof the environmental parameters. Principal coordinate analysis and an analysisusing the Ellenberg ecological indicator values suggested that no importantgradients had been overlooked. Dispersal ability (estimated from dispersalmode)and clumping independent of environmental heterogeneity were related. Poordispersers exhibited stronger clumping at the 100 m scale thangooddispersers. Our results support the joint importance of environmentalheterogeneity and dispersal processes in shaping the mesoscale distributionpatterns of forest herbs. We conclude that the distribution of plant species,and species coexistence and species composition in plant communities, cannot beexplained without simultaneously considering environmental heterogeneity anddispersal.     

Landscape- and field-scale control of spatial variation of soil properties in Mediterranean montane meadows

Soil properties of terrestrial ecosystems are controlled by a variety of factors that operate at different scales. We tested the role of abiotic and biotic factors that potentially influence spatial gradients of total ion content, acidity, carbon, total nitrogen, and total phosphorous in topsoil. We studied a network of Mediterranean montane meadows that spans a 2000-m altitudinal gradient. The analyzed factors were grouped into two spatial scales: a landscape scale (climate and land form) and a field scale (topography, soil texture, soil moisture, and plant community composition). Total ion content and acidity are the major and independent variation trends of soil geochemistry. Soil acidity, carbon, and nitrogen increased along the altitudinal gradient whereas there was no relationship between total ion content and phosphorous and elevation. Climate had no direct influence on the analyzed gradients; all effects of climate were indirect through plant community composition and/or soil moisture. The results point to three types of models that explain the gradients of soil chemical composition: (1) a predominantly biotic control of carbon and nitrogen, (2) a predominantly abiotic control of acidity, and (3) a combined biotic and abiotic control of total ionic content. No direct or indirect effects explained the gradient of phosphorous. In our study region (central Spain), climate is predicted to turn more arid and soils will lose moisture. According to our models, this will result in less acid and fertile soils, and any change in plant community composition will modify gradients of soil carbon, nitrogen, total ion content, and acidity.     

Trait-based community assembly of understory palms along a soil nutrient gradient in a lower montane tropical forest

Two opposing niche processes have been shown to shape the relationship between ecological traits and species distribution patterns: habitat filtering and competitive exclusion. Habitat filtering is expected to select for similar traits among coexisting species that share similar habitat conditions, whereas competitive exclusion is expected to limit the ecological similarity of coexisting species leading to trait differentiation. Here, we explore how functional traits vary among 19 understory palm species that differ in their distribution across a gradient of soil resource availability in lower montane forest in western Panama. We found evidence that habitat filtering influences species distribution patterns and shifts community-wide and intraspecific trait values. Differences in trait values among sites were more strongly related to soil nutrient availability than to variation in light or rainfall. Soil nutrient availability explained a significant amount of variation in site mean trait values for 4 of 15 functional traits. Site mean values of leaf nitrogen and phosphorus increased 37 and 64%, respectively, leaf carbon:nitrogen decreased 38%, and specific leaf area increased 29% with increasing soil nutrient availability. For Geonoma cuneata, the only species occurring at all sites, leaf phosphorus increased 34% and nitrogen:phosphorus decreased 42% with increasing soil nutrients. In addition to among-site variation, most morphological and leaf nutrient traits differed among coexisting species within sites, suggesting these traits may be important for niche differentiation. Hence, a combination of habitat filtering due to turnover in species composition and intraspecific variation along a soil nutrient gradient and site-specific niche differentiation among co-occurring species influences understory palm community structure in this lower montane forest.     

The combined role of topography and overstorey tree composition in promoting edaphic and floristic variation in a Mediterranean forest

This study aimed to address which factors, other than topography, contribute to the floristic variation of forested slopes. The natural forest studied is located in the Sudoeste Alentejano e Costa Vicentina Nature Park (southwestern Portugal). We sampled topographic, edaphic, floristic and community structure variables along three bottom–top hillside transects. Multivariate analyses of soil variables (by PCA), and of woody species composition and floristic–environmental relationships (by CCA and pCCA) were performed. Environmental–floristic trends strongly associated with the elevation gradient were identified. At lower altitudes, the lowest species richness, the highest soil fertility, and the tallest and most dense (least available light) canopy occurred. The spatial variation in woody species composition and abundance was closely associated with Zn availability in the soil and litter groundcover, but these varibles had significant spatial structure in the studied forest. The non-spatially structured species variance was better predicted by soil NO₃^– and NH₄⁺. The spatial variation of species data not shared with environmental variables was also calculated. We suggest that the influence of topographic gradient on the variation of edaphic variables and on the distribution and abundance of woody species was mediated by overstorey tree composition. Locally dominant tree species, in particular Quercus faginea and Quercus suber, may function as ecosystem engineers promoting environmental changes (i.e., Zn availability in soil, litter accumulation and light availability) that influence overall floristic variation.     

Why do mountains support so many species of birds?

Although topographic complexity is often associated with high bird diversity at broad geographic scales, little is known about the relative contributions of geomorphologic heterogeneity and altitudinal climatic gradients found in mountains. We analysed the birds in the western mountains of the New World to examine the two‐fold effect of topography on species richness patterns, using two grains at the intercontinental extent and within temperate and tropical latitudes. Birds were also classified as montane or lowland, based on their overall distributions in the hemisphere. We estimated range in temperature within each cell and the standard deviation in elevation (topographic roughness) based on all pixels within each cell. We used path analysis to test for the independent effects of topographic roughness and temperature range on species richness while controlling for the collinearity between topographic variables. At the intercontinental extent, actual evapotranspiration (AET) was the primary driver of species richness patterns of all species taken together and of lowland species considered separately. In contrast, within‐cell temperature gradients strongly influenced the richness of montane species. Regional partitioning of the data also suggested that range in temperature either by itself or acting in combination with AET had the strongest “effect” on montane bird species richness everywhere. Topographic roughness had weaker “effects” on richness variation throughout, although its positive relationship with richness increased slightly in the tropics. We conclude that bird diversity gradients in mountains primarily reflect local climatic gradients. Widespread (lowland) species and narrow‐ranged (montane) species respond similarly to changes in the environment, differing only in that the richness of lowland species correlates better with broad‐scale climatic effects (AET), whereas mesoscale climatic variation accounts for richness patterns of montane species. Thus, latitudinal and altitudinal gradients in species richness can be explained through similar climatic‐based processes, as has long been argued.     

Phylogenetic community structure in Minnesota oak savanna is influenced by spatial extent and environmental variation

The relative importance of environmental filtering, biotic interactions and neutral processes in community assembly remains an openly debated question and one that is increasingly addressed using phylogenetic approaches. Closely related species may occur together more frequently than expected (phylogenetic clustering) if environmental filtering operates on traits with significant phylogenetic signal. Recent studies show that phylogenetic clustering tends to increase with spatial scale, presumably because greater environmental variation is encompassed at larger spatial scales, providing opportunities for species to sort across environmental gradients. However, if environmental filtering is the cause of species sorting along environmental gradients, then environmental variation rather than spatial scale per se should drive the processes governing community assembly. Using species abundance and light availability data from a long‐term experiment in Minnesota oak savanna understory communities, we explicitly test the hypothesis that greater environmental variation results in greater phylogenetic clustering when spatial scale is held constant. Concordant with previous studies, we found that phylogenetic community structure varied with spatial extent. At the landscape scale (~1000 ha), communities were phylogenetically clustered. At the local scale (0.375ha), phylogenetic community structure varied among plots. As hypothesized, plots encompassing the greatest environmental variation in light availability exhibited the strongest phylogenetic clustering. We also found strong correlations between species functional traits, particularly specific leaf area (SLA) and perimeter per area (PA), and species light availability niche. There was also a phylogenetic signal in both functional traits and species light availability niche, providing a mechanistic explanation for phylogenetic clustering in relation to light availability. We conclude that the pattern of increased phylogenetic clustering with increased environmental variation is a consequence of environmental filtering acting on phylogenetically conserved functional traits. These results indicate that the importance of environmental filtering in community assembly depends not on spatial scale per se, but on the steepness of the environmental gradient.