The assembly of forest communities according to maximum species height along resource and disturbance gradients

An ongoing debate in ecology concerns the relative importance of competition in driving community patterns, especially along gradients of disturbance and productivity. We used a null model to address this question by testing for non‐random organization of forest species according to maximum height, a trait associated with competitive ability for light. Specifically, we compared the species present in 409 different temperate forest plots with the pool of potentially present species (n=639 species), spanning a 50 000 km² area of southern Ontario, Canada. In contrast to current theory, coexisting forest species were neither more similar (i.e. convergent), nor more different (i.e. limiting similarity) in maximum height than expected by chance. However, coexisting forest species had larger maximum heights on average, and their maximum heights were more symmetrically distributed than expected by chance, suggesting that competition has reduced the representation of smaller plant species within plots (i.e. higher turn‐over of species with smaller maximum heights among forest plots). We explored the possibility that our findings resulted from smaller species having relatively narrower niches; however, a conclusive test of this explanation will require knowledge of fundamental, rather than realized niche breadth. We also tested the prediction that the influence of competition changes along gradients of productivity and disturbance by examining how the effect size of our null model tests changed along these gradients. We observed that species with smaller maximum heights were increasingly under‐represented in more productive forest communities, suggesting an increased role for competition in determining species membership in more productive communities. In contrast to theory, however, the effect size of our tests did not significantly change along a gradient of forest disturbance. In summary, we found evidence that maximum species height plays a significant role in driving the non‐random organization of plant species among hundreds of mature forest plots, and that this role is more pronounced in more productive forest plots. 1     

Soil phosphorus and disturbance influence liana communities in a subtropical montane forest

Questions: What are the effects of soil, topography, treefall gaps, tree species composition, and tree density on liana species composition and total liana abundance? Location: A 6‐ha permanent plot in a subtropical montane forest in northwest Argentina. Methods: Multiple regressions were used to quantify associations of liana species composition and total liana abundance with edaphic, disturbance and tree community variables. Gradients in liana and tree species composition were quantified using principal components analysis (PCA). Results: Liana species composition was correlated most strongly with soil phosphorus concentration (R²=0.55). Total liana aanased with phosphorus and the density of recent treefall gaps (R²=0.60). Conclusions: In our study area, liana composition and abundance are most strongly correlated with features of the physical environment, rather than host tree characteristics. Our results support the hypothesis that recent increases in liana abundance in mature tropical forests may be related to increased rates of gap formation.     

Benthic diatom communities in boreal streams: community structure in relation to environmental and spatial gradients

An important goal for community ecology is the characterization and prediction of changes in community patterns along environmental gradients. We aimed to identify the major environmental correlates of diatom distribution patterns in boreal running waters. We classified 197 stream sites based on their diatom flora. Direct ordination methods were then used to identify the key environmental determinants of this diatom-based stream typology. Finally, we tested whether a regional classification scheme based on terrestrial landscapes (ecoregions) provides a reasonable framework for a regional grouping of streams based on their diatom flora. Two-way indicator species analysis produced 13 site groups, which were primarily separated by chemical variables, mainly conductivity, total P and water colour. In partial CCA, the environmental and spatial factors accounted for 38% and 24%, respectively, of explained variation in community composition. A high proportion (almost 40%) of variation explained by the combined effect (spatially-structured environmental) indicated that diatom communities of boreal streams incorporate a strong spatial component. At the level of subecoregions, classification strength was almost equally strong for all sites as for near-pristine reference sites only. Procrustes analysis indicated that spatial factors and patterns in diatom community structure were strongly concordant. Our data support the argument that diatom communities are strongly spatially structured, with distinctly different communities in different parts of the country. Because of the strong spatial patterns of community composition, bioassessment programs utilising lotic diatoms would clearly benefit from regional stratification. A combination of regional stratification and the prediction of assemblage structure from local environmental features might provide the most robust framework for diatom-based assessment of the biological integrity of boreal streams.     

Multivariate analysis of plant communities in the Narok district, Kenya: The influence of environmental factors and human disturbance

Species type in 42 samples was examined using ordination analysis to establish whether the vegetation pattern displays gradation due to environmental factors, and whether the pattern was being altered by human disturbance. The analysis displayed two patterns. First, the ordination on axis 1 demonstrated vegetation gradation from xeric communities on the plains to humid forests on the highland. This pattern is explained on the basis of moisture and nutrient variations. Vegetation vectors based on species presence-absence were correlated with average rainfall, carbon, nitrogen and zinc.The second was separation of disturbed and undisturbed forest and bushland site categories on axis 2 of the ordination space. This pattern suggests that disturbance response is interactive with moisture so that only in moist communities does a significant vegetation response to disturbance occur.     

Effects of environmental and spatial gradients on Quercus-dominated Mountain forest communities in the Hindu-Kush ranges of Pakistan

Quercus-dominated forests are among the most important broad-leaved evergreen forests of the Hindu Kush ranges and are currently prone to drastic anthropogenic and climatic changes. The aim of this study was to provide basic data for the development of a regional oak forest ecosystem framework for ecological restoration and management plan development to maintain local peoples’ livelihoods. Hence, we analyzed distribution patterns and environmental factors that affect regional oak forests’ species composition and diversity. Ward’s Agglomerative clustering divided oak-dominated forest communities into three groups: i.e., Group I, dominated by Quercus baloot had an importance value index (IVI) of 89.87 ± 4.31, Group II, dominated by Quercus dilatata had an IVI of 32.16 ± 15.01, and Group III, dominated by Quercus oblongata had an IVI of 83.14 ± 4.67, respectively. The environmental factors which vary significantly within these communities were latitude, elevation, clay content and bulk density of the soil. Wilting point, saturation point, and electrical conductivity were also considered as ecosystem structural variables. Canonical correspondence analysis (CCA) indicated that community structure was affected by various environmental factors including precipitation, slope angle, elevation, clay content, and relative humidity.     

贵州百里杜鹃风景区旅游干扰对杜鹃群落特征的影响

为揭示旅游干扰对于百里杜鹃风景区杜鹃群落特征影响,选取杜鹃纯林和杜鹃混交林两种杜鹃群落,分析干扰对群落结构及物种多样性的影响。采用典型抽样的方法进行样地设置和调查,通过旅游影响系数分析样地旅游干扰等级,并分析对比了不同旅游干扰程度和干扰方式对群落特征的影响。研究表明：1）百里杜鹃景区共有植被67科102属130种,其中杜鹃纯林共有植被97种,以马缨杜鹃、箭竹和露珠杜鹃为优势物种,杜鹃混交林共有植被115种,以马缨杜鹃、露珠杜鹃和映山红为优势种;2）两种杜鹃群落均在中等干扰水平时具有最高的Berger-Parker丰富度指数和Shannon多样性指数,中度干扰一定程度上促进了杜鹃群落物种多样性增加,不同的是两种群落的Shannon指数变化趋势不同,在杜鹃纯林中表现为中度干扰>轻度干扰>重度干扰,而在杜鹃混交林中中度干扰>重度干扰>轻度干扰,相较于杜鹃纯林,杜鹃混交林物种更为丰富,在旅游干扰下生物多样性更为稳定,抗干扰能力更强,而杜鹃纯林受旅游干扰波动显著;3）垃圾指数、踩踏指数与杜鹃混交林物种多样性指数、群落生长结构无明显相关性,而垃圾指数与杜鹃纯林Shannon多样性指数、物种数以及草本均高呈显著负相关,踩踏指数与杜鹃纯林灌木层高度、盖度呈现极显著负相关,与草本均高呈显著正相关;4）各杜鹃群落对旅游干扰有不同响应,为保持杜鹃群落稳定性,应在花季适当分流,严格控制杜鹃纯林的重度干扰。     

Effects of rain forest disturbance and fragmentation: comparative changes of the raptor community along natural and human-made gradients in French Guiana

A density index of every diurnal raptor species (Falconiformes) was obtained on 101 400 ha sample plots distributed among eight natural habitats and five man-made habitats arranged along gradients of increasing forest degradation and fragmentation. The most significant structural parameter affecting species distribution was the tall canopy forest cover. Species richness, diversity and density all decreased with this mature forest cover index. Individual species and overall community densities decreased along the deforestation gradient but the species richness was partly maintained by species turnover. Six groups of species were identified according to their natural habitat preferences. Their distribution along the deforestation gradient was correlated with their natural habitat selection pattern. Thus the community composition of each vegetation or landscape type was predictable. Fifty-six percent of the regional assemblage of species had their optimal density in the primary forest. A third of them were interior forest species highly sensitive to forest disturbance and opening. The other two-thirds were upper canopy, gap or edge species more tolerant to forest fragmentation. The last twenty-one species were associated with various coastal habitats, from dense forest patches to mangrove and savanna. Again, one third of them were strictly restricted to their specialized habitats while the last two-thirds colonized human-altered habitats and progressively replaced primary forest species with increasing deforestation. The maintenance of large areas of every natural habitat was essential for the conservation of (1) the whole population of a third of the total raptor diversity and (2) optimal and presumably potential source populations of most other species surviving in human-modified habitats.     

Relationships of bird community structure and species distributions to two environmental gradients in the northern boreal forest

Few studies have explicitly examined avian community structure in the North American northern boreal forest Herein we report upon the results of bird surveys in mature stands of boreal forest in the Great Clay Belt Ontario. Canada We related trends in avian community structure and individual bird species abundance to two environmental gradients described in a landbase classification scheme called the Forest Ecosystem Classification a moisture and a nutrient-richness gradient Variation in environmental characteristics is limited in the Ontario Clay Belt and this was reflected in short environmental gradient lengths However major trends in avian community structure were strongly associated with the nutrient-richness gradient axis summarized by this scheme Analyses of avian community composition indicated a continuum from moist coniferous habitats to drier aspen-dominated mixed woodlands with several bird species occurring in varying abundances across the width of both gradients 48 of the 58 species examined showed statistically significant associations with at least one of the two multivariate gradient axes A smaller proportion of short-distance migrant species were associated with these gradients than were species in either the neotropical migrant or resident categories We used a multivariate variable (habitat breadth) to compare degree of habitat specialization across different migratory groups A large proportion of neotropical migrants showed a high degree of habitat specialization m mature Clay Belt forests, and neotropical species with small habitat breadths were more commonly associated with habitats dominated by broad-leaved deciduous tree species than either short-distance or permanent resident species We discuss our findings in relation to the post-glacial history of the Clay Belt region     

Elevational and phytophysiognomic gradients influence the epiphytic community in a cloud forest of the Atlantic phytogeographic domain

Biological communities vary in composition and structure according to changes in the environment, such as in mountainous areas where temperature and moisture change with elevation, leading to gradual changes in vegetation. The altitudinal gradient is considered mirrors of the latitudinal gradient, although the richness peak can occur in the middle of the gradient, due to the greater total availability of moisture and/or due to the mid-domain effect. We aimed to test whether there was an intermediary peak in the richness of vascular epiphytes in two environmental gradients represented by elevation (around 1200–1700 m.s.m.) and by three subphysiognomies of cloud dwarf-forest in the Brazilian Atlantic Forest. For this purpose, 24 plots of 10?×?20 m were established and divided into eight sets in each subphysiognomy, along the altitudinal gradient. The community structure was analyzed by calculating the Shannon diversity index (H′), the Pielou uniformity index (J), and taxonomic diversity indices (Δ⁺ and Λ⁺). In total, 568 phorophytes were sampled, resulting in 3771 occurrences of epiphytes, distributed in 146 species. The diversity was different between the physiognomies of dwarf-forest and along the elevation gradient, and was specifically greater with higher elevation. The value of Δ⁺ was lower than expected in one plot, and represented a potential human impact. The obtained results highlight the diversity of this region and the important contribution of the epiphytes, as well as the sensitivity of epiphytic communities to the different vegetation physiognomies and the climatic variations caused by the elevation.     

Both traits and phylogenetic history influence community structure in snakes over steep environmental gradients

Assemblages of closely related organisms are generated on axes of deep time diversification, biogeographic processes related to dispersal and habitat filtering, and competition. Using models that account for phylogeny, ecology, and traits, we examine how the interaction among biogeography, habitat filtering, and trait convergence influences community assemblage in Nearctic snakes. With 122 community surveys, environmental niche, trait data including size, diet, parity and habitat preference, and a nearly complete phylogeny of snakes from the United States, we ask 1) do phylogenetic species variability (PSV) and traits change in predictable and correlated ways given ecology and geographic distance, 2) are the measured traits variable within and across communities and how is this related to PSV at local scales, and 3) is there evidence of habitat filtering or trait divergence? Following a general trend of western to eastern North American origin and dispersal of major groups, we similarly show a significant decrease in PSV in this direction but unexpectedly with stable trait variance, showing that traits and phylogenetic variability are disconnected at the community level. We also demonstrate that trait variability and not PSV dominates local communities. Finally, regardless of phylogeny, we show that certain traits, such as reproductive mode (parity) frequency, change within communities in response to steep ecological gradients.     

Interactive community responses to disturbance in streams: disturbance history moderates the influence of disturbance types

Both disturbance history and disturbance type act to structure communities through selecting for particular species traits but they may also interact. For example, flooding selects for species with flood‐resistant traits in streams, but those traits could make communities susceptible to other disturbances and so could cause shifts in community composition due to anthropogenic climate change. To better understand the interactive influences of disturbance history and type on community composition, we investigated the response of macroinvertebrate communities to disturbance using in‐stream channels. Using a split‐plot design, individual channels in five ‘stable’ streams and five ‘frequently disturbed’ streams (disturbance history) were subject to different disturbance type treatments (flooding, drying and a control). Disturbance type independently drove effects on species diversity, but all other effects of disturbance type depended on disturbance history. In particular, the interaction of disturbance type and history determined overall community response. Both disturbance types tested produced similar community responses in frequently disturbed streams, including changes in community composition and alterations to the abundance of less mobile taxa, but low‐flow had a significantly greater effect in stable streams. Macroinvertebrate drift was greatest in the rock‐rolling treatments and significantly less in the low‐flow treatment for both disturbance histories. Therefore, disturbance history moderated the effects of disturbance type and determined the mechanism of community response by determining how well species were adapted to disturbance. This outcome suggests that previous disturbances strongly influence how vulnerable communities are to changes in disturbance, and so should be considered when predicting how changes in disturbance regimes will affect future community composition.     

Tropical forest diversity,environmental change and species augmentation: After the intermediate disturbance hypothesis

Abstract. It is not simple to predict how environmental changes may impact tropical forest species diversity. Published hypotheses are almost invariably too incomplete, too poorly specified and too dependent upon unrealistic assumptions to be useful. Ecologists have sought theoretical simplicity, and while this has provided many elegant abstract concepts, it has hindered the attainment of more practical goals. The problem is not how to judge the individual hypotheses and arguments, but rather how to build upon and combine the many hard-won facts and principles into an integrated science. Controversy is inevitable when the assumptions, definitions and applications of a given hypothesis are unclear. Elegance, as an end in itself, has too often been used to justify abstract simplification and a lack of operational definition. Clarifying and combining hypotheses while avoiding assumptions provides a potentially more useful, if less elegant, standpoint. An appraisal of Connell's intermediate disturbance hypothesis, and its application to long-term observations from a Ugandan forest illustrates these concerns. Current emphases encourage ecologists to exclude consideration of environmental instability and non-pristine ecosystems. In reality, many environmental changes and ecological processes contribute to both the accumulation and erosion of diversity, at all spatial and temporal scales. Site histories, contexts, long-term processes, species-pool dynamics, and the role of people require greater emphasis. These considerations reveal that many environmental changes, even those associated with degradation, can lead to a transient rise in species densities. Drawing on related studies, such as forest yield prediction, suggests that the formulation and calibration of simulation models provides the most tractable means to address the complexity of real vegetation. Simulation-based approaches will become increasingly useful both in unifying the study of vegetation dynamics and in providing improved predictive capacity. Quantification of the processes, scales and sensitivities of the dynamics of tropical forest communities remains a major challenge.     

Distribution of pteridophyte communities along environmental gradients in Central Amazonia, Brazil

Extrapolation of local abundance–environment relationships to broader scales provides species distribution models used for conservation planning. We investigated the importance of environmental heterogeneity and geographic distance on pteridophyte species spatial distribution on 38 plots of 250 × 2.5 m distributed over 90 km² in Central Amazon. Inclusion of canopy openness in our models increased the capacity of predicting community composition even under the narrow range of canopy openness found in our plots. Nevertheless, there was still a large amount of unexplained variance (55–65%). The response of the community to the light gradient was hierarchical and we did not find evidences of light partitioning. Most species were concentrated in low light plots but a few common and abundant occurred along the entire gradient. Soil properties were the major determinants of community composition. Contrary to similar studies, slope was not a good predictor of pteridophyte community composition, indicating that this relationship may be site-specific. There was no correlation between floristic distances and geographic distances. We concluded that mesoscale turnover is low, although locally environmental variation determines high turnover of species. Studies among different Amazonian physiognomies tend to find high levels of beta-diversity. However, coarse comparisons can not reveal subtle patterns that are relevant for biodiversity conservation planning. This study found some important changes on pteridophyte community within the same type of forest, mainly related to environmental heterogeneity, even in narrow ranges of environmental variation.     

Environmental gradients and community attributes underlying biodiversity patterns of semi-arid Mediterranean grasslands

Protection and/or establishment of forest plantation have been used as a management strategy to conserve and stop the deterioration of semi-arid Mediterranean grasslands ecosystems, producing a mosaic of vegetation types. This study was intended to investigate the changes in grassland community in response to protection and forest tree plantation practice as well as to explore the underlying environmental gradients responsible for the observed differences or similarities among these vegetation types. Two multivariate analysis methods including discriminate analysis and non-metric multi-dimensional scaling were used to quantify changes in community composition and attributes following different management practices (free grazing, protection with open grassland, sparse and dense forest tree plantations). This was investigated using species frequency, species abundance, or habitat characteristics. The study results showed that habitat types differed significantly between each other and were significantly separated using multivariate approaches. Discrimination based on habitat characteristics and species composition indicated that protection (or grazing) and light (or shade) explained more than 90% of the observed variability in community changes in response to the protection and forest tree plantation. Also, results indicated that shade effect can be attributed to tree canopy cover and/or litter accumulation on the ground. It could be hypothesized that protection from grazing and afforestation resulted in complex environmental gradients of which shade, litter accumulation as well as protection from grazing disturbance are major constituents. A careful manipulation of protection and afforestation can be used to create a spatially different environmental gradients leading to greater habitat diversity as well as a greater species diversity, and better conservation means of grassland in semi-arid areas.     

Tropical montane forest ecotones: climate gradients, natural disturbance, and vegetation zonation in the Cordillera Central, Dominican Republic

Aim We examined relationships between climate–disturbance gradients and patterns of vegetation zonation and ecotones on a subtropical mountain range. Location The study was conducted on the windward slopes of the Cordillera Central, Dominican Republic, where cloud forest appears to shift in a narrow ecotone to monodominant forest of Pinus occidentalis. Methods Climate, disturbance and vegetation data were collected over the elevation range 1100–3100 m and in 50 paired plots along the ecotone. Aerial photographs were georeferenced to a high‐resolution digital elevation model in order to enable the analysis of landscape‐scale patterns of the ecotone. Results A Shipley–Keddy test detected discrete compositional ecotones at 2200 and 2500 m; the distributions of tree species at lower elevations were continuous. The elevation of the ecotone determined with aerial photographs was fairly consistent, namely ± 164 m (SD) over its 124‐km length, but it exhibited significant landscape variation, occurring at a lower elevation in a partially leeward, western zone. The ecotone also occurred significantly lower on ridges than it did in drainage gullies. Ecotone forest structure and composition differed markedly between paired plots. In pine paired plots, the canopy height was 1.7 times higher and the basal area of non‐pine species was 6 times lower than in the cloud forest directly below. Fire evidence was ubiquitous in the pine forest but rare in the abutting cloud forest. Mesoclimate changed discontinuously around the elevation of the ecotone: humidity and cloud formation decreased markedly, and frost frequency increased exponentially. Main conclusions The discreteness of the ecotone was produced primarily by fire. The elevational consistency of the ecotone, however, resulted from the overarching influence of mesoclimate on the elevational patterns of fire occurrence. Declining temperature and precipitation combine with the trade‐wind inversion to create a narrow zone where high‐elevation fires extinguish, enabling fire‐sensitive and fire‐tolerant taxa to abut. Once established, mesotopography and contrasting vegetation physiognomy probably reinforce this boundary through feedbacks on microenvironment and fire likelihood. The prominence of the pine in this study – and of temperate and fire‐tolerant taxa in subtropical montane forests in general – highlights the importance of climate‐disturbance–biogeography interactions in ecotone formation, particularly where fire mediates a dynamic between climate and vegetation.     

海南岛3个林区热带云雾林植物多样性变化

以分布在海南岛西部(霸王岭国家级自然保护区, 21个样方)、西南部(尖峰岭国家级自然保护区, 12个样方)和中部(黎母山省级自然保护区, 15个样方)的热带云雾林为研究对象, 研究α及β物种多样性、功能多样性、谱系多样性的变化, 为植物多样性的保护提供科学依据。结果表明: 尖峰岭群落树木个体多度、物种丰富度最大, 黎母山群落树木个体多度、物种丰富度最小; 黎母山群落间物种组成差异最大, 霸王岭群落间物种组成差异最小, 海南岛霸王岭(西部)、尖峰岭(西南部)和黎母山(中部) 3个林区热带云雾林物种多样性差异可能与空气温度和相对湿度有关。尖峰岭群落内功能丰富度、Rao’s二次熵最低, 功能均匀度最高, 群落间平均成对性状距离最小, 反映群落构建主要受环境筛影响; 霸王岭群落Rao’s二次熵最高, 功能均匀度最低, 群落间平均成对性状距离最大, 反映群落构建主要受限制相似性影响; 黎母山群落内功能丰富度最高, 群落间平均最近性状距离最大, 表明限制相似性在黎母山热带云雾林群落构建中的作用更重要。霸王岭群落内谱系多样性、物种间平均最近相邻谱系距离均较大, 反映物种间谱系关系趋于发散; 黎母山群落内谱系多样性、物种间平均成对谱系距离及群落间平均成对谱系距离均最小, 反映物种间谱系关系趋于聚集; 而尖峰岭群落内物种间平均成对谱系距离、群落间平均成对谱系距离最大, 但物种间平均最近相邻谱系距离最小, 反映物种间谱系关系既具有发散又具有聚集的共存格局。因此, 海南岛热带云雾林群落植物多样性变化格局与环境及物种间相互作用有关。