Methods matter: Different biodiversity survey methodologies identify contrasting biodiversity patterns in a human modified rainforest — A case study with amphibians

Understanding how well tropical forest biodiversity can recover following habitat change is often difficult due to conflicting assessments arising from different studies. One often overlooked potentially confounding factor that may influence assessments of biodiversity response to habitat change, is the possibility that different survey methodologies, targeting the same indicator taxon, may identify different patterns and so lead to different conclusions. Here we investigated whether two different but commonly used survey methodologies used to assess amphibian communities, pitfall trapping and nocturnal transects, indicate the same or different responses of amphibian biodiversity to historic human induced habitat change. We did so in a regenerating rainforest study site located in one of the world’s most biodiverse and important conservation areas: the Manu Biosphere Reserve. We show that the two survey methodologies tested identified contrasting biodiversity patterns in a human modified rainforest. Nocturnal transect surveys indicated biodiversity differences between forest with different human disturbance histories, whereas pitfall trap surveys suggested no differences between forest disturbance types, except for community composition. This pattern was true for species richness, diversity, overall abundance and community evenness and structure. For some fine scale metrics, such as species specific responses and abundances of family groups, both methods detected differences between disturbance types. However, the direction of differences was inconsistent between methods. We highlight that for assessments of rainforest recovery following disturbance, survey methods do matter and that different biodiversity survey methods can identify contrasting patterns in response to different types of historic disturbance. Our results contribute to a growing body of evidence that arboreal species might be more sensitive indicators than terrestrial communities.     

Vertical distribution of epiphytic bryophytes in an Indonesian rainforest

We studied species richness, composition and vertical distribution of epiphytic bryophytes in submontane rainforest of Central Sulawesi. Bryophytes were sampled on eight canopy trees and on eight trees in the forest understorey. Microclimate was measured at trunk bases and at crown bases. The total recorded number of 146 epiphytic bryophyte species is among the highest ever reported for tropical forests and underlines the importance of the Malesian region as a global biodiversity hotspot. Species composition differed significantly between understorey trees and canopy tree trunks on the one hand, and the forest canopy on the other. Fourty-five percent of the bryophyte species were restricted to canopy tree crowns, 12% to the understorey. Dendroid and fan-like species mainly occurred in the forest understorey whereas tufts were most species rich in the tree crowns. The findings reflect the different microclimatic regimes and substrates found in the understorey and in the forest canopy. The results indicate that assessments of the bryophyte diversity of tropical forests are inadequate when understorey trees and tree crowns are excluded.     

Beetle assemblages from an Australian tropical rainforest show that the canopy and the ground strata contribute equally to biodiversity

There remains great uncertainty about how much tropical forest canopies contribute to global species richness estimates and the relative specialization of insect species to vertical zones. To investigate these issues, we conducted a four-year sampling program in lowland tropical rainforest in North Queensland, Australia. Beetles were sampled using a trap that combines Malaise and flight interception trap (FIT) functions. Pairs of this trap, one on the ground and a second suspended 15-20 m above in the canopy were located at five sites, spaced 50 m or more apart. These traps produced 29986 beetles of 1473 species and 77 families. There were similar numbers of individuals (canopy 14473; ground 15513) and species (canopy 1158; ground 895) in each stratum, but significantly more rare species in the canopy (canopy 509; ground 283). Seventy two percent of the species (excluding rare species) were found in both strata. Using IndVal, we found 24 and 27% of the abundant species (n>or=20 individuals) to be specialized to the canopy and the ground strata, respectively, and equivalent analyses at the family level showed figures of 30 and 22%, respectively. These results show that the canopy and the ground strata both provide important contributions to rainforest biodiversity.     

Carbon pools recover more quickly than plant biodiversity in tropical secondary forests

Although increasing efforts are being made to restore tropical forests, little information is available regarding the time scales required for carbon and plant biodiversity to recover to the values associated with undisturbed forests. To address this knowledge gap, we carried out a meta-analysis comparing data from more than 600 secondary tropical forest sites with nearby undisturbed reference forests. Above-ground biomass approached equivalence to reference values within 80 years since last disturbance, whereas below-ground biomass took longer to recover. Soil carbon content showed little relationship with time since disturbance. Tree species richness recovered after about 50 years. By contrast, epiphyte richness did not reach equivalence to undisturbed forests. The proportion of undisturbed forest trees and epiphyte species found in secondary forests was low and changed little over time. Our results indicate that carbon pools and biodiversity show different recovery rates under passive, secondary succession and that colonization by undisturbed forest plant species is slow. Initiatives such as the Convention on Biological Diversity and REDD+ should therefore encourage active management to help to achieve their aims of restoring both carbon and biodiversity in tropical forests.     

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.     

Forest fragmentation and selective logging have inconsistent effects on multiple animal-mediated ecosystem processes in a tropical forest

Forest fragmentation and selective logging are two main drivers of global environmental change and modify biodiversity and environmental conditions in many tropical forests. The consequences of these changes for the functioning of tropical forest ecosystems have rarely been explored in a comprehensive approach. In a Kenyan rainforest, we studied six animal-mediated ecosystem processes and recorded species richness and community composition of all animal taxa involved in these processes. We used linear models and a formal meta-analysis to test whether forest fragmentation and selective logging affected ecosystem processes and biodiversity and used structural equation models to disentangle direct from biodiversity-related indirect effects of human disturbance on multiple ecosystem processes. Fragmentation increased decomposition and reduced antbird predation, while selective logging consistently increased pollination, seed dispersal and army-ant raiding. Fragmentation modified species richness or community composition of five taxa, whereas selective logging did not affect any component of biodiversity. Changes in the abundance of functionally important species were related to lower predation by antbirds and higher decomposition rates in small forest fragments. The positive effects of selective logging on bee pollination, bird seed dispersal and army-ant raiding were direct, i.e. not related to changes in biodiversity, and were probably due to behavioural changes of these highly mobile animal taxa. We conclude that animal-mediated ecosystem processes respond in distinct ways to different types of human disturbance in Kakamega Forest. Our findings suggest that forest fragmentation affects ecosystem processes indirectly by changes in biodiversity, whereas selective logging influences processes directly by modifying local environmental conditions and resource distributions. The positive to neutral effects of selective logging on ecosystem processes show that the functionality of tropical forests can be maintained in moderately disturbed forest fragments. Conservation concepts for tropical forests should thus include not only remaining pristine forests but also functionally viable forest remnants.     

Arthropod Distribution in a Tropical Rainforest: Tackling a Four Dimensional Puzzle

Quantifying the spatio-temporal distribution of arthropods in tropical rainforests represents a first step towards scrutinizing the global distribution of biodiversity on Earth. To date most studies have focused on narrow taxonomic groups or lack a design that allows partitioning of the components of diversity. Here, we consider an exceptionally large dataset (113,952 individuals representing 5,858 species), obtained from the San Lorenzo forest in Panama, where the phylogenetic breadth of arthropod taxa was surveyed using 14 protocols targeting the soil, litter, understory, lower and upper canopy habitats, replicated across seasons in 2003 and 2004. This dataset is used to explore the relative influence of horizontal, vertical and seasonal drivers of arthropod distribution in this forest. We considered arthropod abundance, observed and estimated species richness, additive decomposition of species richness, multiplicative partitioning of species diversity, variation in species composition, species turnover and guild structure as components of diversity. At the scale of our study (2km of distance, 40m in height and 400 days), the effects related to the vertical and seasonal dimensions were most important. Most adult arthropods were collected from the soil/litter or the upper canopy and species richness was highest in the canopy. We compared the distribution of arthropods and trees within our study system. Effects related to the seasonal dimension were stronger for arthropods than for trees. We conclude that: (1) models of beta diversity developed for tropical trees are unlikely to be applicable to tropical arthropods; (2) it is imperative that estimates of global biodiversity derived from mass collecting of arthropods in tropical rainforests embrace the strong vertical and seasonal partitioning observed here; and (3) given the high species turnover observed between seasons, global climate change may have severe consequences for rainforest arthropods.     

Logging impacts on avian species richness and composition differ across latitudes and foraging and breeding habitat preferences

Understanding the causes underlying changes in species diversity is a fundamental pursuit of ecology. Animal species richness and composition often change with decreased forest structural complexity associated with logging. Yet differences in latitude and forest type may strongly influence how species diversity responds to logging. We performed a meta‐analysis of logging effects on local species richness and composition of birds across the world and assessed responses by different guilds (nesting strata, foraging strata, diet, and body size). This approach allowed identification of species attributes that might underlie responses to this anthropogenic disturbance. We only examined studies that allowed forests to regrow naturally following logging, and accounted for logging intensity, spatial extent, successional regrowth after logging, and the change in species composition expected due to random assembly from regional species pools. Selective logging in the tropics and clearcut logging in temperate latitudes caused loss of species from nearly all forest strata (ground to canopy), leading to substantial declines in species richness (up to 27% of species). Few species were lost or gained following any intensity of logging in lower‐latitude temperate forests, but the relative abundances of these species changed substantially. Selective logging at higher‐temperate latitudes generally replaced late‐successional specialists with early‐successional specialists, leading to no net changes in species richness but large changes in species composition. Removing less basal area during logging mitigated the loss of avian species from all forests and, in some cases, increased diversity in temperate forests. This meta‐analysis provides insights into the important role of habitat specialization in determining differential responses of animal communities to logging across tropical and temperate latitudes.     

Spatiotemporal patterns and dynamics of species richness and abundance of woody plant functional groups in a tropical forest landscape of Hainan Island, South China

Tropical forests are among the most species-diverse ecosystems on Earth. Their structures and ecological functions are complex to understand. Functional group is defined as a group of species that play similar roles in an ecosystem. The functional group approach has been regarded as an effective way of linking the compositions of complex ecosystems with their ecological functions. To understand the variation of functional groups in species-rich ecosystems after disturbance, the present study investigated the spatial pattern and temporal dynamics of woody plants in a typically fragmented natural forest landscape of Hainan Island in South China. The study area was classified into eight landscape types based on vegetation type, disturbance manner and the time of recovery. The woody plant species were aggregated into seven functional groups based on the growth form, successional status and plant size. The results gained from the present study showed that all functional groups, except for the emergent and canopy tree species, were present in all eight landscape types. Each landscape type had different numbers of dominant functional groups. There are similar species richness and stem abundance structure among functional groups between mid-successional clear cut lowland rainforest and old growth tropical coniferous forest. This similarity exists in selective logged lowland rainforest and old-growth lowland rainforest, as well as among landscape types of montane rainforest. The functional groups with the same successional status had similar patterns of species richness and stem abundance ratios among different landscape types. The variation patterns of functional groups along the successional stages in terms of species richness and stem abundance among the tropical lowland rainforest landscape types were more similar to each other than those in the tropical montane reinforest landscape types. This study provides further support for the competition-colonization tradeoff and successional niche theory as opposed to models of neutrality and ecological equivalence.     

Biodiversity of leaf-litter ants in fragmented tropical rainforests of Borneo: the value of publically and privately managed forest fragments

In view of the rapid rate of expansion of agriculture in tropical regions, attention has focused on the potential for privately-managed rainforest patches within agricultural land to contribute to biodiversity conservation. However, these sites generally differ in their history of forest disturbance and management compared with other forest fragments, and more information is required on the biodiversity value of these privately-managed sites, particularly in oil-palm dominated landscapes of SE Asia. Here we address this issue, using tropical leaf-litter ants in rainforest fragments surrounded by mature oil palm plantations in Sabah, Borneo as a model system. We compare the species richness and composition of ant assemblages in privately-managed forest fragments (‘high conservation value’ fragments; HCVs) with those in publically-managed fragments of forest (virgin jungle reserves; VJRs) and control sites in extensive tracts of primary forest. In this way, we test the hypothesis that privately-managed and publically-managed forest fragments differ in their species richness and composition as a result of differences in history and management and hence in habitat quality. In support of this hypothesis, we found that HCVs had much poorer habitat quality than VJRs, including lower sizes and densities of trees, less canopy cover, fewer dipterocarp trees and shallower leaf litter. Consequently, HCVs supported only half the species richness of ants in VJRs, which in turn supported 70 % of the species richness of control sites, with vegetation structure and composition explaining 77 % of the variation among forest fragments in ant species richness. HCVs were also much smaller than VJRs but there was only a weak relationship between fragment size and habitat quality, and species richness was not related to fragment size. VJRs supported 78 % of the 156 species found in extensive tracts of forest whereas HCVs supported only 22 %, which was only slightly higher than the proportion previously recorded in oil palm (19 %). These data support previous findings that publically-managed VJR fragments can make an important contribution to biodiversity conservation within agricultural landscapes. However, we suggest that for these HCVs to be effective as reservoirs of biodiversity, management is required to restore vegetation structure and habitat quality, for instance through enrichment planting with native tree species.     

Rainforest seed rain into abandoned tropical Australian pasture is dependent on adjacent rainforest structure and extent

It is well known that the recovery of abandoned tropical pastures to secondary rainforest benefits from the arrival of seeds from adjacent rainforest patches. Less is known, however, about how the structural attributes of adjacent rainforest (e.g. tree density, canopy cover and tree height) impact seed rain patterns into abandoned pastures. Between 2011 and 2013, we used seed traps and ground seed surveys to track the richness and abundance of rainforest seeds entering abandoned pastures in Australia's wet tropics. We also tested how seed rain diversity is related to the distance from forest, the proportion of forest cover in the landscape and several structural attributes of adjacent forest patches, specifically average tree height, canopy cover, tree species richness and density. Almost no seeds were captured in elevated pasture seed traps, even near forest remnants. Abundant forest seeds were found in ground surveys but only within 10 m of forest edges. In ground surveys, seeds from wind‐dispersed species were more abundant, but less species rich, than animal‐dispersed species. A survey of pasture seedling recruits suggested that some forest seeds must be dispersing more than 10 m into pasture at very low frequencies, but only a few species are establishing there. Recruits were predominantly animal‐dispersed not wind‐dispersed species. In addition to distance from forest and the proportion of forest within a 100‐ to 200‐m radius of sampling sites, the richness and density of adjacent forest trees were the most important factors for explaining the probability of seed occurrence in abandoned pastures. Results suggest that without some restoration assistance, the recovery of abandoned pastures into secondary rainforest in Australia's tropical rainforests will likely be limited, at least in part, by a very low rate of seed dispersal away from forest edges and by the diversity and density of trees in adjacent remnant forests.     

Rapid recovery of tropical forest diversity and structure after shifting cultivation in the Philippines uplands

Shifting cultivation is a widespread land‐use in the tropics that is considered a major threat to rainforest diversity and structure. In the Philippines, a country with rich biodiversity and high rates of species endemism, shifting cultivation, locally termed as kaingin, is a major land‐use and has been for centuries. Despite the potential impact of shifting cultivation on forests and its importance to many people, it is not clear how biodiversity and forest structure recover after kaingin abandonment in the country, and how well these post‐kaingin secondary forests can complement the old‐growth forests. We investigated parameters of forest diversity and structure along a fallow age gradient in secondary forests regenerating after kaingin abandonment in Leyte Island, the Philippines (elevation range: 445–650 m asl). We first measured the tree diversity and forest structure indices in regenerating secondary forests and old‐growth forest. We then measured the recovery of tree diversity and forest structure parameters in relation to the old‐growth forest. Finally, using linear mixed effect models (LMM), we assessed the effect of different environmental variables on the recovery of forest diversity and structure. We found significantly higher species density in the oldest fallow sites, while Shannon’s index, species evenness, stem number, basal area, and leaf area index were higher in the old‐growth forest. A homogeneous species composition was found across the sites of older fallow age. Multivariate analysis revealed patch size as a strong predictor of tree diversity and forest structure recovery after shifting cultivation. Our study suggests that, secondary forests regenerating after shifting cultivation abandonment can recover rapidly. Although recovery of forest structure was not as rapid as the tree diversity, our older fallow sites contained a similar number of species as the old‐growth forest. Many of these species are also endemic to the Philippines. Novel and emerging ecosystems like tropical secondary forests are of high conservation importance and can act as a refuge for dwindling tropical forest biodiversity.     

The contribution of cacao agroforests to the conservation of lower canopy ant and beetle diversity in Indonesia

The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.     

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.     

Vertical stratification of seed-dispersing vertebrate communities and their interactions with plants in tropical forests

Vertical stratification (VS) is a widespread phenomenon in plant and animal communities in forests and a key factor for structuring their species richness and biodiversity, particularly in tropical forests. The organisms composing forest communities adjust and shape the complex three-dimensional structure of their environment and inhabit a large variety of niches along the vertical gradient of the forest. Even though the degree of VS varies among different vertebrate groups, patterns of compositional stratification can be observed across taxa. Communities of birds, bats, primates, and non-flying small mammals are vertically stratified in terms of abundance, species richness, diversity, and community composition. Frugivorous members of these taxa play important roles as seed dispersers and forage on fruit resources that, in turn, vary in quantity and nutritional value along the vertical gradient. As a consequence, plant–seed disperser interaction networks differ among strata, which is manifested in differences in interaction frequencies and the degree of mutual specialization. In general, the canopy stratum is composed of strong links and generalized associations, while the lower strata comprise weaker links and more specialized interactions. Investigating the VS of communities can provide us with a better understanding of species habitat restrictions, resource use, spatial movement, and species interactions. Especially in the face of global change, this knowledge will be important as these characteristics can imply different responses of species and taxa at a fine spatial scale.     

Forest disturbance and recovery in Peruvian Amazonia

Amazonian forests function as biomass and biodiversity reservoirs, contributing to climate change mitigation. While they continuously experience disturbance, the effect that disturbances have on biomass and biodiversity over time has not yet been assessed at a large scale. Here, we evaluate the degree of recent forest disturbance in Peruvian Amazonia and the effects that disturbance, environmental conditions and human use have on biomass and biodiversity in disturbed forests. We integrate tree-level data on aboveground biomass (AGB) and species richness from 1840 forest plots from Peru's National Forest Inventory with remotely sensed monitoring of forest change dynamics, based on disturbances detected from Landsat-derived Normalized Difference Moisture Index time series. Our results show a clear negative effect of disturbance intensity tree species richness. This effect was also observed on AGB and species richness recovery values towards undisturbed levels, as well as on the recovery of species composition towards undisturbed levels. Time since disturbance had a larger effect on AGB than on species richness. While time since disturbance has a positive effect on AGB, unexpectedly we found a small negative effect of time since disturbance on species richness. We estimate that roughly 15% of Peruvian Amazonian forests have experienced disturbance at least once since 1984, and that, following disturbance, have been increasing in AGB at a rate of 4.7 Mg ha⁻¹ year⁻¹ during the first 20 years. Furthermore, the positive effect of surrounding forest cover was evident for both AGB and its recovery towards undisturbed levels, as well as for species richness. There was a negative effect of forest accessibility on the recovery of species composition towards undisturbed levels. Moving forward, we recommend that forest-based climate change mitigation endeavours consider forest disturbance through the integration of forest inventory data with remote sensing methods.     

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.     

Functional diversity of epiphytes in two tropical lowland rainforests, French Guiana: using bryophyte life-forms to detect areas of high biodiversity

Recent studies have described a new tropical lowland forest type in the Guianas, the tropical lowland cloud forest. It is characterized by an enriched epiphytic species diversity particularly for bryophytes compared to common lowland rainforest, and is facilitated by frequent early morning fog events in valley locations. While the increase in epiphytic species diversity in lowland cloud forests has been documented, uncertainties remain as to (1) how this small scale variation in water supply is shaping the functional diversity of epiphytic components in lowland forests, and (2) whether information on functional group composition of epiphytes might aid in discerning these cloud forests from the common lowland rainforest. We compare the distribution of functional groups of epiphytes across height zones in lowland cloud forest and lowland rain forest of French Guiana in terms of biomass, cover as well as the composition of bryophyte life-forms. Both forests differed in functional composition of epiphytes in the canopy, in particular in the mid and outer canopy, with the cloud forest having a higher biomass and cover of bryophytes and vascular epiphytes as well as a richer bryophyte life-form composition. Bryophyte life-forms characteristic for cloud forests such as tail, weft and pendants were almost lacking in the canopies of common rain forest whereas they were frequent in lowland cloud forests. We suggest that ground-based evaluation of bryophyte life-form composition is a straightforward approach for identifying lowland cloud forest areas for conservation, which represent biodiversity hotspots in tropical lowland forests.     

Cross-Scale Responses of Biodiversity to Hurricane and Anthropogenic Disturbance in a Tropical Forest

Abstract In studies of biodiversity, considerations of scale—the spatial or temporal domain to which data provide inference—are important because of the non-arithmetic manner in which species richness increases with area (and total abundance) and because fine-scale mechanisms (for example, recruitment, growth, and mortality of species) can interact with broad scale patterns (for example, habitat patch configuration) to influence dynamics in space and time. The key to understanding these dynamics is to consider patterns of environmental heterogeneity, including patterns produced by natural and anthropogenic disturbance. We studied how spatial variation in three aspects of biodiversity of terrestrial gastropods (species richness, species diversity, and nestedness) on the 16-ha Luquillo Forest Dynamics Plot (LFDP) in a tropical forest of Puerto Rico was affected by disturbance caused by Hurricanes Hugo and Georges, as well as by patterns of historic land use. Hurricane-induced changes in spatial organization of species richness differed from those for species diversity. The gamma components of species richness changed after the hurricanes and were significantly different between Hurricanes Hugo and Georges. Alpha and two beta components of species richness, one related to turnover among sites within areas of similar land use and one related to variation among areas of different land use, varied randomly over time after both hurricanes. In contrast, gamma components of species diversity decreased in indistinguishable manners after both hurricanes, whereas the rates of change in the alpha component of species diversity differed between hurricanes. Beta components of diversity related to turnover among sites declined after both hurricanes in a consistent fashion. Those related to turnover among areas with different historic land uses varied stochastically. The immediate effect of hurricanes was to reduce nestedness of gastropod assemblages. Thereafter, nestedness increased during post-hurricane secondary succession, and did so in the same way, regardless of patterns of historic land use. The rates of change in degree of nestedness during secondary succession were different after each hurricane as a result of differences in the severity and extent of the hurricane-induced damage. Our analyses quantified temporal changes in the spatial organization of biodiversity of gastropod assemblages during forest recovery from hurricane-induced damage in areas that had experienced different patterns of historic human land use, and documented the dependence of biodiversity on spatial scale. We hypothesize that cross-scale interactions, likely those between the local demographics of species at the fine scale and the landscape configuration of patches at the broad scale, play a dominant role in affecting critical transfer processes, such as dispersal, and its interrelationship with aspects of biodiversity. Cross-scale interactions have significant implications for the conservation of biodiversity, as the greatest threats to biodiversity arise from habitat modification and fragmentation associated with disturbance arising from human activities.