Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Community structures of Mesostigmata, Prostigmata and Oribatida in broad-leaved regeneration forests and conifer plantations of various ages

The community structures of Mesostigmata, Prostigmata, and Oribatida in the soil of broad-leaved regeneration forests and conifer plantations of various ages were assessed alongside soil and plant environmental variables using three response metrics (density, species richness, and species–abundance distribution). The density and species richness of mites recovered swiftly after clear-cutting or replanting. Oribatid mites dominated the soil mite communities in terms of densities and species richness for both forest types. Soil mite communities in broad-leaved forests was related to forest age, the crown tree communities index, and forest-floor litter weight. In contrast, soil mite communities in the conifer plantation sites were related to various indices of understory plants. The development of the understory plants was synchronized with the silvicultural schedules, including a closed canopy and thinning. Such a conifer plantation management may affect indirectly the community of mites.     

The Effects of Cropping Systems on Avian Communities in Cacao and Banana Agro‐Forestry Systems of Talamanca,Costa Rica

Multispecies agro‐forestry is generally lauded for providing ecosystem services, especially in tropical environments. Avian communities contribute to services such as biodiversity and pest management. Characterizing and evaluating avian community composition in similar cropping systems will help optimize management for ecosystem services. We examined the relationship between cropping system vegetation and avian communities in four shaded agro‐forestry systems common to the Limón province of Costa Rica: abandoned and managed systems of cacao, cacao with banana, and banana. During two field seasons, we detected 2605 birds from 106 species and identified 2791 trees and shrubs from 62 morphospecies. We compared vegetation and avian species richness across systems with mixed‐effects linear models. Canopy, understory, and groundcover vegetation differed among agro‐forestry systems. More ground‐ and understory‐foraging forest species were detected in agro‐forestry systems lacking banana, whereas richness of agricultural generalist species was highest in systems with banana. Richness of understory‐ and ground‐foraging species correlated with understory tree species richness and leaf litter. Our results indicate that shaded cacao and banana systems can have similar canopy‐foraging species richness that includes both agricultural and woodland generalist species, but that interspersing banana with cacao can adversely influence understory forest bird community composition. Agro‐forests with diverse understory vegetation support more understory‐foraging bird species that have proven valuable in pest management. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Dominant forest tree mycorrhizal type mediates understory plant invasions

Forest mycorrhizal type mediates nutrient dynamics, which in turn can influence forest community structure and processes. Using forest inventory data, we explored how dominant forest tree mycorrhizal type affects understory plant invasions with consideration of forest structure and soil properties. We found that arbuscular mycorrhizal (AM) dominant forests, which are characterised by thin forest floors and low soil C : N ratio, were invaded to a greater extent by non‐native invasive species than ectomycorrhizal (ECM) dominant forests. Understory native species cover and richness had no strong associations with AM tree dominance. We also found no difference in the mycorrhizal type composition of understory invaders between AM and ECM dominant forests. Our results indicate that dominant forest tree mycorrhizal type is closely linked with understory invasions. The increased invader abundance in AM dominant forests can further facilitate nutrient cycling, leading to the alteration of ecosystem structure and functions.     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

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.     

Fire rather than nitrogen addition affects understory plant communities in the short term in a coniferous‐broadleaf mixed forest

Increasing fire risk and atmospheric nitrogen (N) deposition have the potential to alter plant community structure and composition, with consequent impacts on biodiversity and ecosystem functioning. This study was conducted to examine short‐term responses of understory plant community to burning and N addition in a coniferous‐broadleaved mixed forest of the subtropical‐temperate transition zone in Central China. The experiment used a pair‐nested design, with four treatments (control, burning, N addition, and burning plus N addition) and five replicates. Species richness, cover, and density of woody and herbaceous plants were monitored for 3 years after a low‐severity fire in the spring of 2014. Burning, but not N addition, significantly stimulated the cover (+15.2%, absolute change) and density (+62.8%) of woody species as well as herb richness (+1.2 species/m², absolute change), cover (+25.5%, absolute change), and density (+602.4%) across the seven sampling dates from June 2014 to October 2016. Light availability, soil temperature, and prefire community composition could be primarily responsible for the understory community recovery after the low‐severity fire. The observations suggest that light availability and soil temperature are more important than nutrients in structuring understory plant community in the mixed forest of the subtropical‐temperate transition zone in Central China. Legacy woody and herb species dominated the understory vegetation over the 3 years after fire, indicating strong resistance and resilience of forest understory plant community and biodiversity to abrupt environmental perturbation.     

Influence of light and soil moisture on Sierran mixed-conifer understory communities

Sierra Nevada forests have high understory species richness yet we do not know which site factors influence herb and shrub distribution or abundance. We examined the understory of an old-growth mixed-conifer Sierran forest and its distribution in relation to microsite conditions. The forest has high species richness (98 species sampled), most of which are herbs with sparse cover and relatively equal abundance. Shrub cover is highly concentrated in discrete patches. Using overstory tree cover and microsite environmental conditions, four habitats were identified; tree cluster, partial canopy, gap, and rock/shallow soil. Herb and shrub species were strongly linked with habitats. Soil moisture, litter depth and diffuse light were the most significant environmental gradients influencing understory plant distribution. Herb cover was most strongly influenced by soil moisture. Shrub cover is associated with more diffuse light, less direct light, and sites with lower soil moisture. Herb richness is most affected by conditions which influence soil moisture. Richness is positively correlated with litter depth, and negatively correlated with direct light and shrub cover. Disturbance or management practices which change forest floor conditions, shallow soil moisture and direct light are likely to have the strongest effect on Sierran understory abundance and richness.     

Interactions with successional stage and nutrient status determines the life‐form‐specific effects of increased soil temperature on boreal forest floor vegetation

The boreal forest is one of the largest terrestrial biomes and plays a key role for the global carbon balance and climate. The forest floor vegetation has a strong influence on the carbon and nitrogen cycles of the forests and is sensitive to changes in temperature conditions and nutrient availability. Additionally, the effects of climate warming on forest floor vegetation have been suggested to be moderated by the tree layer. Data on the effects of soil warming on forest floor vegetation from the boreal forest are, however, very scarce. We studied the effects on the forest floor vegetation in a long‐term (18 years) soil warming and fertilization experiment in a Norway spruce stand in northern Sweden. During the first 9 years, warming favored early successional species such as grasses and forbs at the expense of dwarf shrubs and bryophytes in unfertilized stands, while the effects were smaller after fertilization. Hence, warming led to significant changes in species composition and an increase in species richness in the open canopy nutrient limited forest. After another 9 years of warming and increasing tree canopy closure, most of the initial effects had ceased, indicating an interaction between forest succession and warming. The only remaining effect of warming was on the abundance of bryophytes, which contrary to the initial phase was strongly favored by warming. We propose that the suggested moderating effects of the tree layer are specific to plant life‐form and conclude that the successional phase of the forest may have a considerable impact on the effects of climate change on forest floor vegetation and its feedback effects on the carbon and nitrogen cycles, and thus on the climate.     

Impacts of Small‐scale Clearings due to Selective logging on Dung Beetle Communities

Conservation of biodiversity in production forests is crucial for mitigating biodiversity loss in the tropics. The major ecological impacts of selective logging are often the result of small clearings for skid trails, logging roads, log yards, and logging camps; however, their impacts on forest biodiversity have rarely been examined. The purpose of this study was to assess the impacts of these clearings on a forest‐dependent faunal group, dung beetles, and to identify the environmental factors responsible. Abundance and species richness of dung beetles decreased drastically in clearings, but directly increased in forests with the distance from roads/trails; abundance and species richness at 10 m from roads/trails were almost comparable with those detected in further interior forests. Similarly, species composition was significantly different between forests and clearings (except skid trails) but recovered within a short distance from roads/trails. Canopy openness was the most important environmental factor affecting the abundance, and species richness and composition of dung beetles; most dung beetle species were concentrated under closed forest canopy with less than 10 percent of canopy openness, whereas canopy openness ranged from 16 to 53 percent in clearings. Our study demonstrates that even small‐scale, unpaved clearings affect dung beetle communities through increased canopy openness. Although the effective distance was not very large, a considerable portion of logged areas can be affected when road networks are dense therefore minimizing the density of road networks and enhancing canopy recovery after logging are important for retaining biodiversity in tropical production forests.     

Indicators of restoration success in riparian tropical forests using multiple reference ecosystems

Forest restoration by planting trees often accelerates succession, but the trajectories toward reference ecosystems have rarely been evaluated. Using a chronosequence (4–53 years) of 26 riparian forest undergoing restoration in the Brazilian Atlantic Forest, we modeled how the variables representing forest structure, tree species richness and composition, and the proportion of plant functional guilds change through time. We also estimated the time required for these variables to reach different types of reference ecosystems: old‐growth forest (OGF), degraded forest, and secondary forest. Among the attributes which follow a predictable trajectory over time are: the basal area, canopy cover, density and tree species richness, as well as proportions of shade tolerant and slow growing species or individuals. Most of the variation in density of pteridophythes, lianas, shrubs and phorophythes, proportion of animal‐dispersed individuals, rarefied richness and floristic similarity with reference ecosystems remain unexplained. Estimated time to reach the reference ecosystems is, in general, shorter for structural attributes than for species composition or proportion of functional guilds. The length of this time varies among the three types of reference ecosystems for most attributes. For instance, tree species richness and proportion of shade tolerant or slow growing individuals become similar to secondary forests in about 40 years, but is estimated to take 70 years or more to reach the OGF. Of all the variables considered, canopy cover, basal area, density, and richness of the understory—by their ecological relevance and predictability—are recommended as ecological indicators for monitoring tropical forest restoration success.     

Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette river, Oregon

Riparian forest communities dominated by Populus balsamifera ssp. trichocarpa L. (Torr. and Gray ex Hook.) Brayshaw are important contributors to biodiversity in terrestrial and aquatic ecosystems of the Western United States. Species composition along a successional gradient from stand initiation to late-succession of P. balsamifera-dominated riparian forests was investigated along 145 km of the Willamette River, Oregon. There were 151 total species encountered across 28 stands and a mean species richness of 33.3 species per stand. Young stands were dominated by P. balsamifera and Salix tree spp. and opportunistic herbaceous species. Understory trees, shrubs, and herbaceous species as well as late-successional tree species established 12–15 years after stand initiation. Fraxinus latifolia Benth. was the dominant late-successional tree species. Vertical structural diversity, P. balsamifera mean diameter at breast height, large tree biomass, and stand age were strongly correlated with understory species presence and abundance based on non-metric multidimensional scaling (NMS) ordination. There were no young stands on mid and high terraces and this was reflected in geomorphic position being strongly correlated with the stand age gradient. Abundance of Phalaris arundinacea L. an invasive grass species, was also significantly correlated with plant species composition and abundance. This study indicates that Willamette River riparian forests are diverse and therefore important to the biodiversity of the Willamette River valley and that their presence as a mosaic of communities of different successional stages may be threatened by human interventions, including influences exerted by introduced plant species.     

What shapes cerambycid beetle communities in a tropical forest mosaic? Assessing the effects of host tree identity,forest structure,and vertical stratification

Due to anthropogenic activities, tropical rain forests face many challenges in sustaining biodiversity and maintaining global climates. This study explores how forest successional stage, tree composition, and stratum affect communities of saproxylic cerambycid beetles—concealed feeders that play important roles in forest nutrient cycling. Forty trees in five families (Fabaceae, Lecythidaceae, Malvaceae, Moraceae, and Sapotaceae) were sampled in a mosaic of old‐growth and secondary forest on the Osa Peninsula, Costa Rica. Bait branches yielded 3549 cerambycid individuals in 49 species. Species richness was almost identical in old‐growth and secondary forest, and both yielded specialists, but abundance was higher in old‐growth forest. Overall community structure was most strongly influenced by host plant species; within most plant families it was also impacted by forest successional status. Moraceae was the exception, presumably because the focal tree species was abundant in both old‐growth and secondary forest. Several host and old‐growth specialist species reached high densities within patches of old‐growth forest, but seldom colonized apparently suitable trees within secondary forest. This suggests that even small areas of old‐growth forest can act as refuges, but that secondary forest may act as a barrier to dispersal. The vulnerability of specialized saproxylic insects to land use change will be linked to the ability of their preferred hosts to disperse to and persist in successional habitats; rearing studies may provide the most accurate method to monitor community changes over time.     

Vertical stratification of ichneumonid wasp communities: the effects of forest structure and life‐history traits

Parasitoid wasp communities of the canopy of temperate forests are still largely unexplored. Very little is known about the community composition of parasitoids between canopy and understory and how much of this difference is related to forest structure or parasitoid biological strategies. In this study we investigated upon the difference in the community composition of the parasitic wasps Ichneumonidae between canopy and understory in a lowland temperate forest in northern Italy. We used general linear models to test whether parasitic strategy modifies species vertical stratification and the effect of forest structure. We also tested differences in β‐diversity between canopy and understory traps and over time within single forest layers. We found that stand basal area was positively related to species richness, suggesting that the presence of mature trees can influence local wasp diversity, providing a higher number of microhabitats and hosts. The ichneumonid community of the canopy was different from that of the understory, and the β‐diversity analysis showed higher values for the canopy, due to a higher degree of species turnover between traps. In our analyses, the vertical stratification was different between groups of ichneumonids sharing different parasitic strategies. Idiobiont parasitoids of weakly or deeply concealed hosts were more diverse in the understory than in the canopy while parasitoids of spiders were equally distributed between the two layers. Even though the ichneumonid community was not particularly species‐rich in the canopy of the temperate forests, the extension of sampling to that habitat significantly increased the number of species recorded.     

Community structures of N2‐fixing bacteria associated with the phyllosphere of a Holm oak forest and their response to drought

Biological nitrogen (N) fixation is a key pathway in terrestrial ecosystems and is therefore critical for understanding the responses of ecosystems to global environmental changes. The free‐living diazotrophic community is distributed along the canopy‐to‐soil profile, but the ecological significance of epiphyllic N₂ fixers, despite their functional relevance, on plant foliar surfaces remains very poorly understood compared with the N₂‐fixing community in forest litter and soils. We assessed the community structure of N₂ fixers and overall bacteria by genetic fingerprinting (t‐RFLP) to explore the seasonal successional patterns of the microbial community in the natural phyllosphere of a Holm oak (Quercus ilex) forest submitted to 12‐year field experiment of rain exclusion mimicking the conditions of drought projected for the coming decades. Leaves of Holm oak were analysed in different seasons over a period of 1.5 years. The bacterial community of the phyllosphere did not correspond to the surrounding soil biome in the same area. These analyses provided field evidence for the presence of free‐living diazotrophs associated with the tissues of leaves of Holm oak, the dominant tree species of many Mediterranean forests. The results also revealed that the community composition is affected seasonally and inter‐annually by the environment, and that the composition shifts in response to climate change. Drought treatment increased the richness of the epiphyllic microbial community, especially during the summer. These changes were associated with higher C:N ratios of leaves observed in response to drought in semiarid areas. This epiphyllic microbiota that can potentially fix N₂ extends the capacity of plants to adapt to the environment.     

Influence of Post‐Clearing Treatment on the Recovery of Herbaceous Plant Communities in Amazonian Secondary Forests

Secondary forests are an increasingly common feature in tropical landscapes worldwide and understanding their regeneration is necessary to design effective restoration strategies. It has previously been shown that the woody species community in secondary forests can follow different successional pathways according to the nature of past human activities in the area, yet little is known about patterns of herbaceous species diversity in secondary forests with different histories of land use. We compared the diversity and abundance of herbaceous plant communities in two types of Central Amazonian secondary forests—those regenerating on pastures created by felling and burning trees and those where trees were felled only. We also tested if plant density and species richness in secondary forests are related to proximity to primary forest. In comparison with primary forest sites, forests regenerating on non‐burned habitats had lower herbaceous plant density and species richness than those on burned ones. However, species composition and abundance in non‐burned stands were more similar to those of primary forest, whereas several secondary forest specialist species were found in burned stands. In both non‐burned and burned forests, distance from the forest edge was not related to herbaceous density and species richness. Overall, our results suggest that the natural regeneration of herbaceous species in secondary tropical forests is dependent on a site's post‐clearing treatment. We recommend evaluating the land history of a site prior to developing and implementing a restoration strategy, as this will influence the biological template on which restoration efforts are overlaid.     

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.     

Restoration of Tropical Moist Forests on Bauxite-Mined Lands in the Brazilian Amazon

We evaluated forest structure and composition in 9- to 13-year-old stands established on a bauxite-mined site at Trombetas (Pará), Brazil, using four different reforestation techniques following initial site preparation and topsoil replacement. These techniques included reliance on natural forest regeneration, mixed commercial species plantings of mostly exotic timber trees, direct seeding with mostly native early successional tree species, and mixed native species plantings of more than 70 tree species (the current operational restoration treatment at this site). Replicated fixed-radius plots in each treatment and in undisturbed primary forest were used to quantify the canopy and understory structure and the abundance and diversity of all vascular plant species. Treatment comparisons considered regeneration density, species richness and diversity for all floristic categories, and, for trees and shrubs, the relative contribution of initial planting and subsequent regeneration from soil seed banks and seed inputs from nearby primary forests. With the possible exception of the stands of mixed commercial species, which were superior to all others in terms of tree basal-area development but relatively poor in species richness, all treatments were structurally and floristically diverse, with a high probability of long-term restoration success. Of these, the mixed native species plantings appeared to be at least risk of arrested succession due to the dominance of a broader range of tree species of different successional stages or expected life spans. In all treatments, several locally important families of primary forest trees (Annonaceae, Chrysobalanaceae, Lauraceae, Palmae and Sapotaceae) were markedly underrepresented due to a combination of poor survival of initial plantings and limitations on seed dispersal from the surrounding primary forest.     

Resilience of tropical rain forests: tree community reassembly in secondary forests

Understanding the recovery dynamics of ecosystems presents a major challenge in the human-impacted tropics. We tested whether secondary forests follow equilibrium or non-equilibrium dynamics by evaluating community reassembly over time, across different successional stages, and among multiple life stages. Based on long-term and static data from six 1-ha plots in NE Costa Rica, we show that secondary forests are undergoing reassembly of canopy tree and palm species composition through the successful recruitment of seedlings, saplings, and young trees of mature forest species. Such patterns were observed over time within sites and across successional stages. Floristic reassembly in secondary forests showed a clear convergence with mature forest community composition, supporting an equilibrium model. This resilience stems from three key factors co-occurring locally: high abundance of generalist species in the regional flora, high levels of seed dispersal, and local presence of old-growth forest remnants.