The impact of fragmentation and density regulation on forest succession in the Atlantic rain forest

The Atlantic Rain Forest, an important biodiversity hot spot, has faced severe habitat loss since the last century which has resulted in a highly fragmented landscape with a large number of small forest patches (<100 ha). For conservation planning it is essential to understand how current and future forest regeneration depends on ecological processes, fragment size and the connection to the regional seed pool. We have investigated the following questions by applying the forest growth simulation model FORMIND to the situation of the Atlantic Forest in the state of Sao Paulo, SE Brazil: (1) which set of parameters describing the local regeneration and level of density regulation can reproduce the biomass distribution and stem density of an old growth forest in a reserve? (2) Which additional processes apart from those describing the dynamics of an old growth forest, drive forest succession of small isolated fragments? (3) Which role does external seed input play during succession? Therefore, more than 300 tree species have been classified into nine plant functional types (PFTs), which are characterized by maximum potential height and shade tolerance. We differentiate between two seed dispersal modes: (i) local dispersal, i.e. all seedlings originated from fertile trees within the simulated area and (ii) external seed rain. Local seed dispersal has been parameterized following the pattern oriented approach, using biomass estimates of old growth forest. We have found that moderate density regulation is essential to achieve coexistence for a broad range of regeneration parameters. Considering the expected uncertainty and variability in the regeneration processes it is important that the forest dynamics are robust to variations in the regeneration parameters. Furthermore, edge effects such as increased mortality at the border and external seed rain have been necessary to reproduce the patterns for small isolated fragments. Overall, simulated biomass is much lower in the fragments compared to the continuous forest, whereas shade tolerant species are affected most strongly by fragmentation. Our simulations can supplement empirical studies by extrapolating local knowledge on edge effects of fragments to larger temporal and spatial scales. In particular our results show the importance of external seed rain and therefore highlight the importance of structural connectivity between regenerating fragments and mature forest stands. (C) 2009 Elsevier B.V. All rights reserved.

Succession and management of tropical dry forests in the Americas: Review and new perspectives

Understanding tropical forest succession is critical for the development of tropical forest conservation strategies worldwide, given that tropical secondary forests can be considered the forests of the future. Tropical dry forests (TDF) are among the most threatened tropical ecosystems, there are more secondary forests and forest restoration efforts that require a better understanding of successional processes. The main goal of this synthesis for this special issue on the ecology and management of tropical dry forests in the Americas is to present a summarized review of the current knowledge of the ecology and management implications associated to TDF succession. We explore specific issues associated to tropical dry forest succession with emphasis on the use of chronosequences, plant diversity and composition, plant phenology and remote sensing, pollination, and animal-plant interactions; all under the integrating umbrella of ecosystem succession. We also emphasize the need to conduct socio-ecological research to understand changes in land-use history and its effects on succession and forest regeneration of TDF. We close this paper with some thoughts and ideas associated with the strong need for an integrating dimension not considered until today: the role of cyberinfrastructure and eco-informatics as a tool to support sound conservation, management and understanding of TDF in the Americas. (C) 2009 Elsevier B.V. All rights reserved.

Habitat structure and food resources for wildlife across successional stages in a tropical forest

Tropical forests are experiencing an increase in the proportion of secondary forests as a result of the balance between the widespread harvesting of old-growth forests and the regeneration in abandoned areas. The impacts of such a process on biodiversity are poorly known and intensely debated. Recent reviews and multi-taxa studies indicate that species replacement in wildlife assemblages is a consistent pattern, sometimes stronger than changes in diversity, with a replacement from habitat generalists to old-growth specialists being commonly observed during tropical forest regeneration. However, the ecological drivers of such compositional changes are rarely investigated, despite its importance in assessing the conservation value of secondary forests, and to support and guide management techniques for restoration. By sampling 28 sites in a continuous Atlantic forest area in Southeastern Brazil, we assessed how important aspects of habitat structure and food resources for wildlife change across successional stages, and point out hypotheses on the implications of these changes for wildlife recovery. Old-growth areas presented a more complex structure at ground level (deeper leaf litter, and higher woody debris volume) and higher fruit availability from an understorey palm, whereas vegetation connectivity, ground-dwelling arthropod biomass, and total fruit availability were higher in earlier successional stages. From these results we hypothetize that generalist species adapted to fast population growth in resource-rich environments should proliferate and dominate earlier successional stages, while species with higher competitive ability in resource-limited environments, or those that depend on resources such as palm fruits, on higher complexity at the ground level, or on open space for flying, should dominate older-growth forests. Since the identification of the drivers of wildlife recovery is crucial for restoration strategies, it is important that future work test and further develop the proposed hypotheses. We also found structural and functional differences between old-growth forests and secondary forests with more than 80 years of regeneration, suggesting that restoration strategies may be crucial to recover structural and functional aspects expected to be important for wildlife in much altered ecosystems, such as the Brazilian Atlantic forest. (C) 2012 Elsevier B.V. All rights reserved.

Effect of rodents on seed fate of five hornbill-dispersed tree species in a tropical forest in north-east India

Hornbills are important dispersers of a wide range of tree species. Many of these species bear fruits with large, lipid-rich seeds that could attract terrestrial rodents. Rodents have multiple effects on seed fates, many of which remain poorly understood in the Palaeotropics. The role of terrestrial rodents was investigated by tracking seed fate of five horn bill-dispersed tree species in a tropical forest in north-cast India. Seeds were marked inside and outside of exclosures below 6-12 parent fruiting trees (undispersed seed rain) and six hornbill nest trees (a post-dispersal site). Rodent visitors and seed removal ere monitored using camera traps. Our findings suggest that several rodent species. especially two species of porcupine were major on-site seed predators. Scatter-hoarding was rare (1.4%). Seeds at hornbill nest trees had lower survival compared with parent fruiting trees, indicating that clumped dispersal by hornbills may not necessarily improve seed survival. Seed survival in the presence and absence of rodents varied with tree species. Some species (e.g. Polyalthia simiarum) showed no difference, others (e.g. Dysoxylum binectariferum) experienced up to a 64%. decrease in survival in the presence of rodents. The differing magnitude of seed predation by rodents can have significant consequences at the seed establishment stage.

Endophytic fungal communities in woody perennials of three tropical forest types of the Western Ghats, southern India

Fungal endophytes of tropical trees are expected to be exceptionally species rich as a consequence of high tree diversity in the tropics and the purported host restriction among the endophytes. Based on this premise, endophytes have been regarded as a focal group for estimating fungal numbers because their possible hyperdiverse nature would reflect significantly global fungal diversity. We present our consolidated ten-year work on 75 dicotyledonous tree hosts belonging to 33 families and growing in three different types of tropical forests of the NBR in the Western Ghats, southern India. We conclude that endophyte diversity in these forests is limited due to loose host affiliations among endophytes. Some endophytes have a wide host range and colonize taxonomically disparate hosts suggesting adaptations in them to counter a variety of defense chemicals in their hosts. Furthermore, such polyphagous endophytes dominate the endophyte assemblages of different tree hosts. Individual leaves may be densely colonized but only by a few endophyte species. It appears that the environment (the type of forest in this case) has a larger role in determining the endophyte assemblage of a plant host than the taxonomy of the host plant. Thus, different tropical plant communities have to be studied for their endophyte diversity to test the generalization that endophytes are hyperdiverse in the tropics, estimate their true species richness, and use them as a predictor group for more accurate assessment of global fungal diversity.

Averting biodiversity collapse in tropical forest protected areas

The rapid disruption of tropical forests probably imperils global biodiversity more than any other contemporary phenomenon(1-3). With deforestation advancing quickly, protected areas are increasingly becoming final refuges for threatened species and natural ecosystem processes. However, many protected areas in the tropics are themselves vulnerable to human encroachment and other environmental stresses(4-9). As pressures mount, it is vital to know whether existing reserves can sustain their biodiversity. A critical constraint in addressing this question has been that data describing a broad array of biodiversity groups have been unavailable for a sufficiently large and representative sample of reserves. Here we present a uniquely comprehensive data set on changes over the past 20 to 30 years in 31 functional groups of species and 21 potential drivers of environmental change, for 60 protected areas stratified across the world's major tropical regions. Our analysis reveals great variation in reserve `health': about half of all reserves have been effective or performed passably, but the rest are experiencing an erosion of biodiversity that is often alarmingly widespread taxonomically and functionally. Habitat disruption, hunting and forest-product exploitation were the strongest predictors of declining reserve health. Crucially, environmental changes immediately outside reserves seemed nearly as important as those inside in determining their ecological fate, with changes inside reserves strongly mirroring those occurring around them. These findings suggest that tropical protected areas are often intimately linked ecologically to their surrounding habitats, and that a failure to stem broad-scale loss and degradation of such habitats could sharply increase the likelihood of serious biodiversity declines.

Performance of established native seedlings in relation to invasive Lantana camara, rainfall and species' habitat preferences in a seasonally dry tropical forest

Native species' response to the presence of invasive species is context specific. This response cannot be studied in isolation from the prevailing environmental stresses in invaded habitats such as seasonal drought. We investigated the combined effects of an invasive shrub Lantana camara L. (lantana), seasonal rainfall and species' microsite preferences on the growth and survival of 1,105 naturally established seedlings of native trees and shrubs in a seasonally dry tropical forest. Individuals were followed from April 2008 to February 2010, and growth and survival measured in relation to lantana density, seasonality of rainfall and species characteristics in a 50-ha permanent forest plot located in Mudumalai, southern India. We used a mixed effects modelling approach to examine seedling growth and generalized linear models to examine seedling survival. The overall relative height growth rate of established seedlings was found to be very low irrespective of the presence or absence of dense lantana. 22-month growth rate of dry forest species was lower under dense lantana while moist forest species were not affected by the presence of lantana thickets. 4-month growth rates of all species increased with increasing inter-census rainfall. Community results may be influenced by responses of the most abundant species, Catunaregam spinosa, whose growth rates were always lower under dense lantana. Overall seedling survival was high, increased with increasing rainfall and was higher for species with dry forest preference than for species with moist forest preference. The high survival rates of naturally established seedlings combined with their basal sprouting ability in this forest could enable the persistence of woody species in the face of invasive species.

Contrasting fire regimes in a seasonally dry tropical forest and a savanna ecosystem in the western Ghats, India

Tropical dry forests and savannas constitute more than half of all tropical forests and grasslands, but little is known about forest fire regimes within these two extensive types of ecosystems. Forest fire regimes in a predominantly dry forest in India, the Nilgiri landscape, and a predominantly savanna ecosystem in the Sathyamangalam landscape, were examined. Remote sensing data were applied to delineate burned areas, determine fire size characteristics, and to estimate fire-rotation intervals. Belt transects (0.5 ha) were used to estimate forest structure, diversity, and fuel loads. Mean area burned, mean number of fires, and mean fire size per year were substantially higher in the Nilgiri landscape compared to the Sathyamangalam landscape. Mean fire-rotational interval was 7.1 yr in the Nilgiri landscape and 44.1 yr in the Sathyamangalam landscape. Tree (>= 10 cm diameter at breast height) species diversity, tree density, and basal area were significantly higher in the Nilgiri landscape compared to the Sathyamangalam landscape. Total fuel loads were significantly higher in tropical dry and moist deciduous forests in the Nilgiri landscape, but total fuel loads were higher in the tropical dry thorn forests of the Sathyamangalam landscape. Thus, the two landscapes revealed contrasting fire regimes and forest characteristics, with more and four-fold larger fires in the Nilgiri landscape. The dry forests and savannas could be maintained by a combination of factors, such as fire, grazing pressures, and herbivore populations. Understanding the factors maintaining these two ecosystems will be critical for their conservation.

Long-Term Environmental Correlates of Invasion by Lantana camara (Verbenaceae) in a Seasonally Dry Tropical Forest

Invasive species, local plant communities and invaded ecosystems change over space and time. Quantifying this change may lead to a better understanding of the ecology and the effective management of invasive species. We used data on density of the highly invasive shrub Lantana camara (lantana) for the period 1990-2008 from a 50 ha permanent plot in a seasonally dry tropical forest of Mudumalai in southern India. We used a cumulative link mixed-effects regression approach to model the transition of lantana from one qualitative density state to another as a function of biotic factors such as indicators of competition from local species (lantana itself, perennial grasses, invasive Chromolaena odorata, the native shrub Helicteres isora and basal area of native trees) and abiotic factors such as fire frequency, inter-annual variability of rainfall and relative soil moisture. The density of lantana increased substantially during the study period. Lantana density was negatively associated with the density of H. isora, positively associated with basal area of native trees, but not affected by the presence of grasses or other invasive species. In the absence of fire, lantana density increased with increasing rainfall. When fires occurred, transitions to higher densities occurred at low rainfall values. In drier regions, lantana changed from low to high density as rainfall increased while in wetter regions of the plot, lantana persisted in the dense category irrespective of rainfall. Lantana seems to effectively utilize resources distributed in space and time to its advantage, thus outcompeting local species and maintaining a population that is not yet self-limiting. High-risk areas and years could potentially be identified based on inferences from this study for facilitating management of lantana in tropical dry forests.

Characterising weather patterns associated with fire in a seasonally dry tropical forest in southern India

Anthropogenic fires in seasonally dry tropical forests are a regular occurrence during the dry season. Forest managers in India, who presently follow a fire suppression policy in such forests, would benefit from a system of assessing the potential risk to fire on a particular day. We examined the relationship between weather variables (seasonal rainfall, relative humidity, temperature) and days of fire during the dry seasons of 2004-2010, based on MODIS fire incident data in the seasonally dry tropical forests of Mudumalai in the Western Ghats, southern India. Logistic regression analysis showed that high probabilities of a fire day, indicating successful ignition of litter and grass fuel on the forest floor, were associated with low levels of early dry season rainfall, low daily average relative humidity and high daily average temperatures. These weather conditions are representative of low moisture levels of fine fuels, suggesting that the occurrence of fire is moderated by environmental conditions that reduce the flammability of fine fuels in the dry tropics. We propose a quantitative framework for assessing risk of a fire day to assist forest managers in anticipating fire occurrences in this seasonally dry tropical forest, and possibly for those across South Asia.

Reduced Hornbill Abundance Associated with Low Seed Arrival and Altered Recruitment in a Hunted and Logged Tropical Forest

Logging and hunting are two key direct threats to the survival of wildlife in the tropics, and also disrupt important ecosystem processes. We investigated the impacts of these two factors on the different stages of the seed dispersal cycle, including abundance of plants and their dispersers and dispersal of seeds and recruitment, in a tropical forest in north-east India. We focused on hornbills, which are important seed dispersers in these forests, and their food tree species. We compared abundances of hornbill food tree species in a site with high logging and hunting pressures (heavily disturbed) with a site that had no logging and relatively low levels of hunting (less disturbed) to understand logging impacts on hornbill food tree abundance. We compared hornbill abundances across these two sites. We, then, compared the scatter-dispersed seed arrival of five large-seeded tree species and the recruitment of four of those species. Abundances of hornbill food trees that are preferentially targeted by logging were two times higher in the less disturbed site as compared to the heavily disturbed site while that of hornbills was 22 times higher. The arrival of scatter-dispersed seeds was seven times higher in the less disturbed site. Abundances of recruits of two tree species were significantly higher in the less disturbed site. For another species, abundances of younger recruits were significantly lower while that of older recruits were higher in the heavily disturbed site. Our findings suggest that logging reduces food plant abundance for an important frugivore-seed disperser group, while hunting diminishes disperser abundances, with an associated reduction in seed arrival and altered recruitment of animal-dispersed tree species in the disturbed site. Based on our results, we present a conceptual model depicting the relationships and pathways between vertebrate-dispersed trees, their dispersers, and the impacts of hunting and logging on these pathways.

Regeneration of Juvenile Woody Plants after Fire in a Seasonally Dry Tropical Forest of Southern India

Woody tree species in seasonally dry tropical forests are known to have traits that help them to recover from recurring disturbances such as fire. Two such traits are resprouting and rapid post-fire growth. We compared survival and growth rates of regenerating small-sized individuals (juveniles) of woody tree species after dry season fire (February-March) at eight adjacent pairs of burnt and unburnt transects in a seasonally dry tropical forest in southern India. Juveniles were monitored at 3-mo intervals between August 2009 and August 2010. High juvenile survivorship (>95%) was observed in both burnt and unburnt areas. Growth rates of juveniles, analyzed at the community level as well as for a few species individually (especially fast-growing ones), were distinctly higher in burnt areas compared to unburnt areas after a fire event, particularly during the pre-monsoon season immediately after a fire. Rapid growth by juveniles soon after a fire may be due to lowered competition from other vegetative forms such as grasses, possibly aided by the availability of resources stored belowground. Such an adaptation would allow a juvenile bank to be retained in the understory of a dry forest, from where individuals can grow to a possible fire-tolerant size during favorable conditions.