Elevation‐dependent effects of forest fragmentation on plant–bird interaction networks in the tropical Andes

Tropical forests harbor diverse ecological communities of plants and animals that are organized in complex interaction networks. The diversity and structure of plant–animal interaction networks may change along elevational gradients and in response to human‐induced habitat fragmentation. While previous studies have analyzed the effects of elevation and forest fragmentation on species interaction networks in isolation, to our knowledge no study has investigated whether the effects of forest fragmentation on species interactions may differ along elevational gradients. In this study, we analyzed main and interaction effects of elevation and forest fragmentation on plant–frugivore interaction networks at plant and bird species level. Over a period spanning two years, we recorded plant–frugivore interactions at three elevations (1000, 2000 and 3000 m a.s.l.) and in two habitat types (continuous and fragmented forest) in tropical montane forests in southern Ecuador. We found a consistent effect of elevation on the structure of plant–frugivore networks. We observed a decrease in the number of effective bird partners of plants and, thus, a decline in the redundancy of bird species with increasing elevation. Furthermore, bird specialization on specific plant partners increased towards high elevations. Fragmentation had a relatively weak effect on the interaction networks for both plant and bird species, but resulted in a significant increase in bird specialization in fragmented forests at high elevations. Our results indicate that forest fragmentation may have stronger effects on plant–frugivore interaction networks at high compared to low elevations because bird species richness declined more steeply towards high elevations than plant species richness. We conclude that conservation efforts should prioritize the maintenance of consumer diversity, for instance by maintaining stretches of continuous forest. This applies in particular to species‐poor communities, such as those at high elevations, as the ecological processes in these communities seem most sensitive towards forest fragmentation.     

Richness,species composition and functional groups in Agaricomycetes communities along a vegetation and elevational gradient in the Andean Yungas of Argentina

The Neotropics are among the least explored regions from a mycological perspective. A few recent molecular studies in South America have shown high fungal diversity as well as numerous groups of mostly undescribed taxa. Through soil metabarcoding analysis we compared richness and species composition among macrofungal communities, belonging to Agaricales, Russulales, Boletales and Phallomycetidae groups, in three elevational forests types in the subtropical Yungas of Northwestern Argentina (Piedmont forest; Montane forest, Montane cloud forest). The aims of this study were to assess richness of taxonomic and functional groups along the elevation gradient and to assess the relationships between environmental variables and species composition in the studied fungal communities. The results have shown rich Agaricomycetes communities, diversely structured among forests habitats. The elevation gradient differentially affected the richness and distribution of Agaricales, Russulales, Boletales and Phallomycetidae. Based on fungal trophic modes and guilds, the gradient also affected the ectomycorrhizal taxa distribution. When considering the basidiomata growth forms (agaricoid, boletoid, gasteroid, etc.), only the secotioid type showed significant elevational differences. Additional analyses indicated that saprotrophic nutritional mode was dominant along the entire gradient, being partially replaced by biotrophic modes at higher elevations. Fungal communities in the Montane cloud forests are most dissimilar when compared with communities at the Piedmont forest and Montane forest, which is consistent with the different biogeographic origins of these forests. DNA metabarcoding sequence analysis provided detailed information on the diversity and taxonomic and functional composition of macrofungal communities.     

Vertically stratified frugivore community composition and interaction frequency in a liana fruiting across forest strata

Vertical stratification is a key feature of tropical forests and structures plant–frugivore interactions. However, it is unclear whether vertical differences in plant-frugivore interactions are due to differences among strata in plant community composition or inherent preferences of frugivores for specific strata. To test this, we observed fruit removal of a diverse frugivore community on the liana Marcgravia longifolia in a Peruvian rain forest. Unlike most other plants, Marcgravia longifolia produces fruits across forest strata. This enabled us to study effects of vertical stratification on fruit removal without confounding effects of plant species and stratum. We found a high number of visits of a few frugivore species in the understorey and a low number of visits of many different frugivores in the canopy and midstorey. Whereas partial and opportunistic frugivores foraged across strata with differing frequencies, obligate frugivores were only found eating fruits in the higher strata. Avian frugivores foraging in the canopy were mainly large species with pointed wings, whereas under- and midstorey avian foragers were smaller with rounded wings. Our findings suggest a continuous shift in the frugivore community composition along the vertical gradient, from a few generalized frugivores in the understorey to a diverse set of specialized frugivores in the canopy. This shift in the frugivore community leads to correlated, reciprocal changes from specialized to generalized plant-frugivore interactions. Thus, we conclude that vertical niche differentiation between species in tropical forests persists even when food resources are available across strata. This highlights its role for promoting biodiversity and ecosystem functioning.     

Past Human Disturbance Effects upon Biodiversity are Greatest in the Canopy; A Case Study on Rainforest Butterflies

A key part of tropical forest spatial complexity is the vertical stratification of biodiversity, with widely differing communities found in higher rainforest strata compared to terrestrial levels. Despite this, our understanding of how human disturbance may differentially affect biodiversity across vertical strata of tropical forests has been slow to develop. For the first time, how the patterns of current biodiversity vary between three vertical strata within a single forest, subject to three different types of historic anthropogenic disturbance, was directly assessed. In total, 229 species of butterfly were detected, with a total of 5219 individual records. Butterfly species richness, species diversity, abundance and community evenness differed markedly between vertical strata. We show for the first time, for any group of rainforest biodiversity, that different vertical strata within the same rainforest, responded differently in areas with different historic human disturbance. Differences were most notable within the canopy. Regenerating forest following complete clearance had 47% lower canopy species richness than regenerating forest that was once selectively logged, while the reduction in the mid-storey was 33% and at ground level, 30%. These results also show for the first time that even long term regeneration (over the course of 30 years) may be insufficient to erase differences in biodiversity linked to different types of human disturbance. We argue, along with other studies, that ignoring the potential for more pronounced effects of disturbance on canopy fauna, could lead to the underestimation of the effects of habitat disturbance on biodiversity, and thus the overestimation of the conservation value of regenerating forests more generally.     

Tree Species Richness Strengthens Relationships between Ants and the Functional Composition of Spider Assemblages in a Highly Diverse Forest

In species‐rich ecosystems, such as subtropical and tropical forests, higher trophic level interactions are key mediators of ecosystem functioning. Plant species loss may alter these interactions, but the effects of plant diversity might be modified by intraguild interactions, particularly among predators. We analyzed the relationships between spiders and ants, two dominant predatory arthropod taxa, on tree saplings across a gradient from medium to high woody plant species richness in a subtropical forest in Southeast China. Neither ant nor spider total biomass was significantly related to plant species richness. By contrast, the biomass distribution of web‐building and hunting spiders changed and spider family richness increased in the presence of ants, resulting in more web builder‐dominated assemblages. However, these relationships depended on the plant communities, and were stronger in plots with higher plant species richness. Our results indicate that in addition to potential effects of ants on hunting spiders in particular, ants could indirectly influence intraguild interactions within spider assemblages. The observed shifts in the spider assemblages with increasing ant presence and plant species richness may have functional consequences, as web‐building and hunting spiders have distinct prey spectra. The relationships among ants, spiders, and plant species richness might contribute to explaining the non‐significant relationship between the overall effects of predators and plant diversity previously observed in the same forest plots. Our findings thus give insight into the complexity of biotic interactions in such species‐rich ecosystems.     

太白山北坡落叶阔叶林物种多样性特征

对太白山北坡落叶阔叶林从不同类型,层次的丰富度,均匀度和物种多样性指数及其与环境梯度的关系等方面进行了分析。结果表明,处于中山带的锐齿槲栎林及辽东栎林的多样性较高,高海拔与低海拔处的群落多样性较低。水热梯度对物种多样性有很大影响。植物生长型与物种多样性的关系表现为灌木层的多样性指数（Ｄ）和均匀度大于草本层,而草本层大于乔木层,草木层物种丰富度最大,灌木层次之,乔木层最小,不同群落间灌木层各多样性测     

Fungal community composition in neotropical rain forests: the influence of tree diversity and precipitation

Plant diversity is considered one factor structuring soil fungal communities because the diversity of compounds in leaf litter might determine the extent of resource heterogeneity for decomposer communities. Lowland tropical rain forests have the highest plant diversity per area of any biome. Since fungi are responsible for much of the decomposition occurring in forest soils, understanding the factors that structure fungi in tropical forests may provide valuable insight for predicting changes in global carbon and nitrogen fluxes. To test the role of plant diversity in shaping fungal community structure and function, soil (0-20?cm) and leaf litter (O horizons) were collected from six established 1-ha forest census plots across a natural plant diversity gradient on the Isthmus of Panama. We used 454 pyrosequencing and phospholipid fatty acid analysis to evaluate correlations between microbial community composition, precipitation, soil nutrients, and plant richness. In soil, the number of fungal taxa increased significantly with increasing mean annual precipitation, but not with plant richness. There were no correlations between fungal communities in leaf litter and plant diversity or precipitation, and fungal communities were found to be compositionally distinct between soil and leaf litter. To directly test for effects of plant species richness on fungal diversity and function, we experimentally re-created litter diversity gradients in litter bags with 1, 25, and 50 species of litter. After 6?months, we found a significant effect of litter diversity on decomposition rate between one and 25 species of leaf litter. However, fungal richness did not track plant species richness. Although studies in a broader range of sites is required, these results suggest that precipitation may be a more important factor than plant diversity or soil nutrient status in structuring tropical forest soil fungal communities.     

Do foliar fungal communities of Norway spruce shift along a tree species diversity gradient in mature European forests?

Foliar fungal species are diverse and colonize all plants, though whether forest tree species composition influences the distribution of these fungal communities remains unclear. Fungal communities include quiescent taxa and the functionally important and metabolically active taxa that respond to changes in the environment. To determine fungal community shifts along a tree species diversity gradient, needles of Norway spruce were sampled from trees from four mature European forests. We hypothesized that the fungal communities and specific fungal taxa would correlate with tree species diversity. Furthermore, the active fungal community, and not the total community, would shift along the tree diversity gradient. High-throughput sequencing showed significant differences in the fungal communities in the different forests, and in one forest, tree diversity effects were observed, though this was not a general phenomenon. Our study also suggests that studying the metabolically active community may not provide additional information about community composition or diversity.     

Forest plant diversity is threatened by Robinia pseudoacacia (black-locust) invasion

The effects of black-locust invasion on plant forest diversity are still poorly investigated. Vascular plants are likely to be influenced by increasing nutrient availability associated with the nitrogen-fixing activity of black-locust, whereas it is not clear if, along with stand aging, black-locust formations regain forest species. The main aim of the present study was to test whether the increase of black-locust stand age promoted a plant variation in mature stands leading to assemblages similar to those of native forests. Therefore, plant richness and composition of stands dominated by native trees were compared with pure black-locust stands of different successional stages. Our study confirmed that the replacement of native forests by pure black-locust stands causes both plant richness loss and shifts in species composition. In black-locust stands plant communities are dominated by nitrophilous species and lack many of the oligothrophic and acidophilus species typical of native forests. Plant communities of native forests are more diverse with respect to pure black-locust stands, suggesting that black-locust invasion also causes a homogenization of the plant forest biota. We did not detect differences across the successional gradient of black-locust stands, and mature stands do not recover the diversity of plant species which are lost by the replacement of the native forests by black-locust. Accordingly some efforts in reducing the negative impacts of black-locust invasion on plant forest biota should be focused at least in those areas where conservation is among management priorities, such in the case of habitats included in the Habitat Directive (92/43 ECE).     

Environmental correlates of plant diversity in Korean temperate forests

Mountainous areas of the Korean Peninsula are among the biodiversity hotspots of the world's temperate forests. Understanding patterns in spatial distribution of their species richness requires explicit consideration of different environmental drivers and their effects on functionally differing components. In this study, we assess the impact of both geographical and soil variables on the fine-scale (400 m²) pattern of plant diversity using field data from six national parks, spanning a 1300 m altitudinal gradient. Species richness and the slopes of species–area curves were calculated separately for the tree, shrub and herb layer and used as response variables in regression tree analyses. A cluster analysis distinguished three dominant forest communities with specific patterns in the diversity–environment relationship. The most widespread middle-altitude oak forests had the highest tree richness but the lowest richness of herbaceous plants due to a dense bamboo understory. Total richness was positively associated with soil reaction and negatively associated with soluble phosphorus and solar radiation (site dryness). Tree richness was associated mainly with soil factors, although trees are frequently assumed to be controlled mainly by factors with large-scale impact. A U-shaped relationship was found between herbaceous plant richness and altitude, caused by a distribution pattern of dwarf bamboo in understory. No correlation between the degree of canopy openness and herb layer richness was detected. Slopes of the species–area curves indicated the various origins of forest communities. Variable diversity–environment responses in different layers and communities reinforce the necessity of context-dependent differentiation for the assessment of impacts of climate and land-use changes in these diverse but intensively exploited regions.     

Relative importance of climate vs local factors in shaping the regional patterns of forest plant richness across northeast China

Northeast (NE) China covers three climatic zones and contains all the major forest types of NE Asia. We sampled 108 forest plots in six nature reserves across NE China to examine the influence of climate and local factors (canopy seasonality, successional stage, topography and forest structure) on geographic patterns of plant richness. We analyzed the relative effects of different factors at two spatial scales: the regional scale (across both latitude and altitude) and the local scale (along the altitudinal gradient within site). Our results showed that the relative importance of climate vs local factors differed remarkably depending on scale and functional group. While total and tree species richness were mainly limited by climate, herb and shrub richness was more related to local factors (especially at the local scale). In the climatic factors, heat sum was the major correlate of tree, shrub and total species richness, while herb richness was more associated with winter coldness. Precipitation was not a limiting factor for forest plant richness in NE China. Climate accounted for 34–76% of variation in richness at the regional scale, but explained only 0–44% at the local scale. Among the local factors, shrub species richness was sensitive to seasonal canopy openness, with higher richness in deciduous forests than in the evergreen needle-leaf forest. On the other hand, herb richness was sensitive to forest successional stage, with higher richness in middle- successional forests than in the early and late-sucessional forests. Local topography (aspect and position on slope) and forest structure (tree density) also showed remarkable influence on species richness. Our results suggest the importance of including local factors when examining large scale diversity gradient (especially for understory species), and the necessity of comparing diversity patterns among functional groups at different spatial scales.     

Bat and bird diversity along independent gradients of latitude and tree composition in European forests

Species assemblages are shaped by local and continental-scale processes that are seldom investigated together, due to the lack of surveys along independent gradients of latitude and habitat types. Our study investigated changes in the effects of forest composition and structure on bat and bird diversity across Europe. We compared the taxonomic and functional diversity of bat and bird assemblages in 209 mature forest plots spread along gradients of forest composition and vertical structure, replicated in 6 regions spanning from the Mediterranean to the boreal biomes. Species richness and functional evenness of both bat and bird communities were affected by the interactions between latitude and forest composition and structure. Bat and bird species richness increased with broadleaved tree cover in temperate and especially in boreal regions but not in the Mediterranean where they increased with conifer abundance. Bat species richness was lower in forests with smaller trees and denser understorey only in northern regions. Bird species richness was not affected by forest structure. Bird functional evenness increased in younger and denser forests. Bat functional evenness was also influenced by interactions between latitude and understorey structure, increasing in temperate forests but decreasing in the Mediterranean. Covariation between bat and bird abundances also shifted across Europe, from negative in southern forests to positive in northern forests. Our results suggest that community assembly processes in bats and birds of European forests are predominantly driven by abundance and accessibility of feeding resources, i.e., insect prey, and their changes across both forest types and latitudes.     

The Importance of Primary Tropical Rain Forest For Species Diversity: An Investigation Using Arboreal Ants as an example

The arboreal ant communities of a primary lowland rain forest and three differently disturbed forests that lie close together forming an anthropogenic disturbance gradient were collected with insecticide fogging. Combined samples from all trees (87 foggings) comprised 153,504 ants sorted to 331 morphospecies. The primary forest ant fauna was characterized by high species richness and 53 foggings were necessary to collect communities representatively. Another 63 species of ants were found in the disturbed forests indicating a large regional species pool that might exceed 420 species of arboreal ants. Anthropogenic disturbance caused a change in the taxonomic composition, diversity and structure of ant communities. Community size was a predictor of species richness in the severely disturbed forest types but not in the old secondary or primary forest. Ant abundance had declined significantly in the disturbed forests and only 10% of the primary forest’s species were collected in the most disturbed forest type. In each of the secondary forests a change in the frequency distribution of species was observed and a small number of species had gained numerical dominance. Analysis of species associations indicates that the strength of species interactions changed with the degree of forest disturbance. These changes were still clearly recognizable after 40 years of forest regeneration despite optimal conditions for colonization from the adjacent primary forest, demonstrating that the time scale needed for forest recovery after anthropogenic disturbance is very long.     

Pattern of local plant species richness along a gradient of landscape topographical heterogeneity: result of spatial mass effect or environmental shift?

Several processes are hypothesised to mediate the relationship between local (microsite) plant species richness and the topographical heterogeneity of the surrounding landscape. In a topographically heterogeneous landscape with various habitats occurring close to each other, local species richness may be enriched by species from surrounding habitats due to the spatial mass effect (sink‐source dynamics). In contrast, increased habitat fragmentation due to spatial heterogeneity may have a negative effect on local species richness. The spatial mass effect is thought to be more pronounced in communities with a higher ratio of generalists, as generalists are more likely to establish viable populations in sink habitats. To reveal the pattern of local species richness along a gradient of landscape topographical heterogeneity at middle altitudes of the Bohemian Massif, we used 2551 forest vegetation plots stored in the Czech National Phytosociological Database. We developed an analytical approach relating the pattern of local species richness of vegetation types to the gradient of landscape topographical heterogeneity. An increase or decrease in species richness with increasing landscape heterogeneity was related to changes in the generalist/specialist ratio, and also to changes in soil reaction and productivity estimated through Ellenberg indicator values. Local species richness along a gradient of increasing landscape heterogeneity increased in nutrient‐poor vegetation and decreased in nutrient‐rich vegetation. Nutrient‐poor vegetation types, such as thermophilous and acidophilous oak forests, also had a high proportion of habitat generalists, supporting the hypothesis that increased richness in heterogeneous landscapes may result from the spatial mass effect. However, the same pattern may be explained by a shift in environmental conditions along the landscape heterogeneity gradient, such as increasing productivity of nutrient‐rich vegetation types or increasing soil reaction of most vegetation types in more heterogeneous landscapes. We discuss available evidence and conclude that these two explanations need not be mutually exclusive.     

Seed dispersal networks in tropical forest fragments: Area effects,remnant species,and interaction diversity

Seed dispersal interactions involve key ecological processes in tropical forests that help to maintain ecosystem functioning. Yet this functionality may be threatened by increasing habitat loss, defaunation, and fragmentation. However, generalist species, and their interactions, can benefit from the habitat change caused by human disturbance while more specialized interactions mostly disappear. Therefore, changes in the structure of the local, within fragment, networks can be expected. Here we investigated how the structure of seed dispersal networks changes along a gradient of increasing habitat fragmentation. We analyzed 16 bird seed dispersal assemblages from forest fragments of a biodiversity-rich ecosystem. We found significant species–, interaction–, and network–area relationships, yet the later was determined by the number of species remaining in each community. The number of frugivorous bird and plant species, their interactions, and the number of links per species decreases as area is lost in the fragmented landscape. In contrast, network nestedness has a negative relationship with fragment area, suggesting an increasing generalization of the network structure in the gradient of fragmentation. Network specialization was not significantly affected by area, indicating that some network properties may be invariant to disturbance. Still, the local extinction of partner species, paralleled by a loss of interactions and specialist–specialist bird–plant seed dispersal associations, suggests the functional homogenization of the system as area is lost. Our study provides empirical evidence for network–area relationships driven by the presence/absence of remnant species and the interactions they perform.     

Butterfly Community Composition Across a Successional Gradient in a Human‐disturbed Afro‐tropical Rain Forest

Knowledge of the recovery of insect communities after forest disturbance in tropical Africa is very limited. Here, fruit‐feeding butterflies in a tropical rain forest at Kibale National Park, Uganda, were used as a model system to uncover how, and how fast, insect communities recover after forest disturbance. We trapped butterflies monthly along a successional gradient for one year. Traps were placed in intact primary forest compartments, heavily logged forest compartments with and without arboricide treatment approximately 43 years ago, and in conifer‐clearcut compartments, ranging from 9 to 19 years of age. The species richness, total abundance, diversity, dominance, and similarity of the community composition of butterflies in the eight compartments were compared with uni‐ and multivariate statistics. A total of 16,728 individuals representing 88 species were trapped during the study. Butterfly species richness, abundance, and diversity did not show an increasing trend along the successional gradient but species richness and abundance peaked at intermediate stages. There was monthly variation in species richness, abundance, diversity and composition. Butterfly community structure differed significantly among the eight successional stages and only a marginal directional change along the successional gradient emerged. The greatest number of indicator species and intact forest interior specialists were found in one of the primary forests. Our results show that forest disturbance has a long‐term impact on the recovery of butterfly species composition, emphasizing the value of intact primary forests for butterfly conservation.     

Importance of the understory stratum to entomofaunal diversity in a temperate deciduous forest

The vertical stratification of lepidopteran and coleopteran communities in a cool-temperate deciduous forest in Japan was examined to evaluate the hypothesis of an expected uniform distribution of mobile flying insects between the canopy and understory of temperate forests. Lepidopteran and coleopteran insects were trapped using light traps at three sites in each of the canopy and understory for three consecutive nights each month from April to October 2001. For Lepidoptera, species richness, abundance, and family richness were significantly higher in the understory than in the canopy. For Coleoptera, only abundance was larger in the canopy relative to the understory; species and family richness did not differ between the strata. The beta diversity of the lepidopteran community was larger between the strata than among sites, but the coleopteran community showed an inverse pattern. These results imply the presence of vertical stratification within the lepidopteran community, but not within the coleopteran community, in the temperate forest. The understory contributes more than the canopy to lepidopteran diversity in the temperate forest, although this stratification may be relatively weak because, in contrast to the situation in tropical forests, the canopy and understory assemblages share many species.     

长白山溪流河岸带森林木本植物多样性沿海拔梯度分布规律

采用样带网格调查方法和α、β多样性指数分析方法,研究了长白山河岸带原始林和次生林群落木本植物多样性沿海拔梯度分布规律及其对采伐干扰的响应.结果表明：不同海拔区域河岸带原始林群落均由11～13个树种组成,其在群落中的地位随海拔升高而发生变化,阔叶树种优势地位逐步被针叶树种所取代,采伐干扰不仅改变了群落树种丰富度及其在群落中的地位,而且使针叶树种取代阔叶树种的趋势有所增强;河岸带原始林群落物种多样性沿海拔梯度呈现出中、低海拔区域相对较高且比较恒定(2.454～2.544),高海拔区域(2.250)下降的分布规律,采伐干扰改变了其沿海拔梯度分布格局(波动型),加大了不同海拔区域群落间的波动性(2.174～2.692);河岸带原始林群落树种沿海拔梯度的变动速率相对较低(1.5～3.5),且群落相似性较高(0.85～0.94),采伐干扰使次生林群落树种沿海拔梯度的变动幅度增大(0.5～6.0),群落相似性下降(0.68～0.91),但次生林群落沿海拔梯度分布仍具有较高连续性.