A Study of Plant Species Extinction in Singapore: Lessons for the Conservation of Tropical Biodiversity

The native vascular plant flora of the Republic of Singapore has suffered the extinction of 594 out of a total 2277 species. These represent local, not global, species extinctions. Coastal habitats, including mangroves, have lost 39% of their species, while inland forests have last 29%. Epiphytic species (62% loss) appear particularly prone to extinction, which is reflected in a similar disposition exhibited by the Orchidaceae. Deforestation and disturbance have been the main cause of plant species extinction in Singapore. The rich mangrove epiphyte flora has been totally exterminated, and a number of tree species are reduced to populations of a few mature individuals. Many more species continue to survive than the species-area relationship would predict given the 99.8% loss of primary forest. This is interpreted as a result of the failure of equilibrium to be achieved yet in the remnant forest fragments, even after more than a century of isolation. Singapore's secondary forests appear to accrete plant diversity very slowly, even if contiguous with primary forest areas. We conclude that remnant fragments of primary tropical forest, even of very small size, can play a major role in the conservation of tropical biodiversity. The patterns of extinction observed in Singapore indicate that coastal and estuarine sites are in greatest demand for development and therefore must be given high priority for conservation despite their somewhat lower biodiversity. Epiphyte and orchid diversity appear to be very good indicators of the degree of disturbance suffered by a habitat in the humid tropics.     

Epiphyte Biodiversity in the Coffee Agricultural Matrix: Canopy Stratification and Distance from Forest Fragments

Abstract: Quality of the agricultural matrix profoundly affects biodiversity and dispersal in agricultural areas. Vegetatively complex coffee agroecosystems maintain species richness at larger distances from the forest. Epiphytes colonize canopy trees and provide resources for birds and insects and thus effects of agricultural production on epiphytes may affect other species. We compared diversity, composition, and vertical stratification of epiphytes in a forest fragment and in two coffee farms differing in management intensity in southern Mexico. We also examined spatial distribution of epiphytes with respect to the forest fragment to examine quality of the two agricultural matrix types for epiphyte conservation. We sampled vascular epiphytes in a forest fragment, a shade polyculture farm, and a shade monoculture farm at 100 m, 200 m, and 400 m from the forest. Epiphyte and orchid richness was greater in the forest than in the monoculture but richness was similar in the forest and polyculture farm. Epiphyte species composition differed with habitat type, but not with distance from the forest. In the forest, epiphytes were distributed throughout tree canopies, but in the farms, epiphytes were primarily found on trunks and larger branches. Epiphyte richness and species similarity to forest species declined with distance from the forest fragment in the monoculture, but richness and similarity to forest species did not decline with distance from forest in the polyculture. This suggests polyculture coffee has greater conservation value. In contrast, monoculture coffee is likely a sink habitat for epiphytes dispersing from forests into coffee. Coffee farms differ from forests in terms of the habitat they provide and species composition, thus protecting forest fragments is essential for epiphyte conservation. Nonetheless, in agricultural landscapes, vegetatively complex coffee farms may contribute to conservation of epiphytes more than other agricultural land uses.     

Assessing the strength of climate and land-use influences on montane epiphyte communities

Epiphytes, air plants that are structurally dependent on trees, are a keystone group in tropical forests; they support the food and habitat needs of animals and influence water and nutrient cycles. They reach peak diversity in humid montane forests. Climate predictions for Central American mountains include increased temperatures, altered precipitation seasonality, and increased cloud base heights, all of which may challenge epiphytes. Although remaining montane forests are highly fragmented, many tropical agricultural systems include trees that host epiphytes, allowing epiphyte communities to persist even in landscapes with lower forest connectivity. I used structural equations models to test the relative effects of climate, land use, tree characteristics, and biotic interactions on vascular epiphyte diversity with data from 31 shade coffee farms and 2 protected forests in northern Nicaragua. I also tested substrate preferences of common species with randomization tests. Tree size, tree diversity, and climate all affected epiphyte richness, but the effect of climate was almost entirely mediated by bryophyte cover. Bryophytes showed strong sensitivity to mean annual temperature and insolation. Many ferns and some orchids were positively associated with bryophyte mats, whereas bromeliads tended to establish among lichen or on bare bark. The tight relationships between bryophytes and climate and between bryophytes and vascular epiphytes indicated that relatively small climate changes could result in rapid, cascading losses of montane epiphyte communities. Currently, shade coffee farms can support high bryophyte cover and diverse vascular epiphyte assemblages when larger, older trees are present. Agroforests serve as valuable reservoirs for epiphyte biodiversity and may be important early-warning systems as the climate changes.     

The Value of Primary, Secondary, and Plantation Forests for a Neotropical Herpetofauna

Abstract:  Plantation forests and second-growth forests are becoming dominant components of many tropical forest landscapes. Yet there is little information available concerning the consequences of different forestry options for biodiversity conservation in the tropics. We sampled the leaf-litter herpetofauna of primary, secondary, and Eucalyptus plantation forests in the Jari River area of northeastern Brazilian Amazonia. We used four complementary sampling techniques, combined samples from 2 consecutive years, and collected 1739 leaf-litter amphibians (23 species) and 1937 lizards (30 species). We analyzed the data for differences among forest types regarding patterns of alpha and beta diversity, species-abundance distributions, and community structure. Primary rainforest harbored significantly more species, but supported a similar abundance of amphibians and lizards compared with adjacent areas of second-growth forest or plantations. Plantation forests were dominated by wide-ranging habitat generalists. Secondary forest faunas contained a number of species characteristic of primary forest habitat. Amphibian communities in secondary forests and Eucalyptus plantations formed a nested subset of primary forest species, whereas the species composition of the lizard community in plantations was distinct, and was dominated by open-area species. Although plantation forests are relatively impoverished, naturally regenerating forests can help mitigate some negative effects of deforestation for herpetofauna. Nevertheless, secondary forest does not provide a substitute for primary forest, and in the absence of further evidence from older successional stands, we caution against the optimistic claim that natural forest regeneration in abandoned lands will provide refuge for the many species that are currently threatened by deforestation .     

A Contemporary Assessment of Change in Humid Tropical Forests

Abstract: In recent decades the rate and geographic extent of land‐use and land‐cover change has increased throughout the world's humid tropical forests. The pan‐tropical geography of forest change is a challenge to assess, and improved estimates of the human footprint in the tropics are critical to understanding potential changes in biodiversity. We combined recently published and new satellite observations, along with images from Google Earth and a literature review, to estimate the contemporary global extent of deforestation, selective logging, and secondary regrowth in humid tropical forests. Roughly 1.4% of the biome was deforested between 2000 and 2005. As of 2005, about half of the humid tropical forest biome contained 50% or less tree cover. Although not directly comparable to deforestation, geographic estimates of selective logging indicate that at least 20% of the humid tropical forest biome was undergoing some level of timber harvesting between 2000 and 2005. Forest recovery estimates are even less certain, but a compilation of available reports suggests that at least 1.2% of the humid tropical forest biome was in some stage of long‐term secondary regrowth in 2000. Nearly 70% of the regrowth reports indicate forest regeneration in hilly, upland, and mountainous environments considered marginal for large‐scale agriculture and ranching. Our estimates of the human footprint are conservative because they do not resolve very small‐scale deforestation, low‐intensity logging, and unreported secondary regrowth, nor do they incorporate other impacts on tropical forest ecosystems, such as fire and hunting. Our results highlight the enormous geographic extent of forest change throughout the humid tropics and the considerable limitations of the science and technology available for such a synthesis.     

Conservation of Vascular Epiphyte Diversity in Mexican Coffee Plantations

Abstract:  Coffee plantations have replaced many lower-montane forests in the Neotropics, and ongoing intensification is converting traditional polycultures with a variety of shade trees to plantations with no or monospecific shade trees. To evaluate the impact of coffee cultivation on epiphyte diversity, I surveyed vascular epiphytes on shade trees in nine different coffee plantations and compared records with those in four natural forests in central Veracruz, Mexico. Eighty-nine species occurred in coffee plantations, and 104 species occurred in natural forests. The number of epiphytic species in traditional polycultures with old shade trees, mostly of the genus Inga , was similar to that in forests, but plantations with small trees and sparse shade hosted fewer epiphytes than those with large trees. Epiphyte communities were, however, more homogeneous in coffee plantations than in forests, possibly because of a drier microclimate and the lack of large and long-lived trees. These results demonstrate the value of traditional polycultures for epiphyte diversity, show the importance of conserving large shade trees, and suggest that these coffee systems may not be suitable for all epiphytes.     

Effects of Forest Fragmentation and Habitat Degradation on West African Leaf-Litter Frogs

Abstract:  Habitat degradation alters the dynamics and composition of anuran assemblages in tropical forests. The effects of forest fragmentation on the composition of anuran assemblages are so far poorly known. We studied the joint influence of forest fragmentation and degradation on leaf-litter frogs. We specifically asked whether the processes structuring leaf-litter anuran assemblages in fragmented forests are the same as those in continuous forests. We analyzed anuran assemblages with respect to habitat characteristics, including fragmentation and degradation parameters. In comparison with continuous forests, species richness and diversity were lower and assemblage composition was altered in forest fragments. These changes seemed to be mainly caused by habitat degradation rather than forest fragmentation. Availability of aquatic sites for breeding, vegetation structure (including those variables indicating degradation), and leaf-litter cover had the most influence on the presence of single species. The comparatively small impact of fragmentation on anurans might be due to the location of the study area; it still possessed large tracts of continuous forest. These forest blocks may stabilize the regional rainforest climate and thus weaken the effects of fragmentation .     

Canopy Seed Banks as Time Capsules of Biodiversity in Pasture-Remnant Tree Crowns

Abstract: Tropical pastures present multiple barriers to tree regeneration and restoration. Relict trees serve as “regeneration foci” because they ameliorate the soil microclimate and serve as safe spots for dispersers. Here, we describe another mechanism by which remnant trees may facilitate pasture regeneration: the presence of seed banks in the canopy soil that accumulates from decomposing epiphytes within the crowns of mature remnant trees in tropical cloud forest pastures. We compared seed banks of canopy soils (histosols derived from fallen leaves, fruits, flower, and twigs of host trees and epiphytes, dead bryophytes, bark, detritus, dead animals, and microorganisms, and dust that accumulate on trunks and the upper surfaces of large branches) in pastures, canopy soils in primary forest trees, and soil on the forest floor in Monteverde, Costa Rica. There were 5211 epiphytic and terrestrial plant seeds in the three habitats. All habitats were dominated by seeds in a relatively small number of plant families, most of which were primarily woody, animal pollinated, and animal dispersed. The density of seeds on the forest floor was greater than seed density in either pasture‐canopy or forest‐canopy soils; the latter two did not differ. Eight species in 44 families and 61 genera from all of the habitats were tallied. There were 37 species in the pasture‐canopy soil, 33 in the forest‐canopy soil, and 57 on the forest floor. Eleven species were common to all habitats. The mean species richness in the pasture canopy was significantly higher than the forest canopy (F =83.38; p < 0.02). Nonmetric multidimensional scaling ordination revealed that the communities were distinct. Greenhouse experiments verified that many of these seeds were viable, with 29 taxa germinating (23 taxa in pruned mats [mimic of exposed conditions] and 16 taxa in control mats [intact conditions]) within 2 months of observation. Nearly half the species that germinated were characteristic of primary forests (primary forest samples, 19%; pasture samples, 29%). This supports the idea that canopy seed banks of pasture trees can function as time capsules by providing propagules that are removed in both space and time from the primary forest. Their presence may enhance the ability of pastures to regenerate more quickly, reinforcing the importance of trees in agricultural settings.     

Effects of Pioneer Tree Species Hyperabundance on Forest Fragments in Northeastern Brazil

Abstract: Despite many studies on fragmentation of tropical forests, the extent to which plant and animal communities are altered in small, isolated forest fragments remains obscure if not controversial. We examined the hypothesis that fragmentation alters the relative abundance of tree species with different vegetative and reproductive traits. In a fragmented landscape (670 km²) of the Atlantic Forest of northeastern Brazil, we categorized 4056 trees of 182 species by leafing pattern, reproductive phenology, and morphology of seeds and fruit. We calculated relative abundance of traits in 50 1‐ha plots in three types of forest configurations: forest edges, small forest fragments (3.4–83.6 ha), and interior of the largest forest fragment (3500 ha, old growth). Although evergreen species were the most abundant across all configurations, forest edges and small fragments had more deciduous and semideciduous species than interior forest. Edges lacked supra‐annual flowering and fruiting species and had more species and stems with drupes and small seeds than small forest fragments and forest interior areas. In an ordination of species similarity and life‐history traits, the three types of configurations formed clearly segregated clusters. Furthermore, the differences in the taxonomic and functional (i.e., trait‐based) composition of tree assemblages we documented were driven primarily by the higher abundance of pioneer species in the forest edge and small forest fragments. Our work provides strong evidence that long‐term transitions in phenology and seed and fruit morphology of tree functional groups are occurring in fragmented tropical forests. Our results also suggest that edge‐induced shifts in tree assemblages of tropical forests can be larger than previously documented.     

Frog, Bat, and Dung Beetle Diversity in the Cloud Forest and Coffee Agroecosystems of Veracruz, Mexico

Abstract:  We compared the species diversity of copronecrophagous beetles (Scarabaeinae) , bats, and frogs in tropical montane cloud forest (original vegetation) and shaded coffee plantations (an agroecosystem common to the region) for a landscape in central Veracruz, Mexico. We sampled in three tropical montane cloud forest fragments and in three coffee plantations with traditional polyculture shade between 1998 and 2001. The three indicator groups responded differently to the transformation of tropical montane cloud forest into shaded coffee plantations. The species richness of frogs was one-fifth less in coffee plantations than in forest fragments, and only one-third of the frog species occurred in both forest fragments and coffee plantations. The number of beetle species and their abundance was significantly greater in coffee plantations than in the forest fragments, whereas species richness and species composition of bats were virtually the same in both habitats. The majority of the abundant species remained as such in both communities, but species that were less abundant were not scarce in both habitats. We attributed differences in the species assemblages to the differing degrees of penetrability of the borders of the two habitat types (especially for the coffee plantations) and to the differences in life-history traits among species. Shaded coffee plantations form a matrix that envelops the remaining fragments of cloud forest. Together they connect the forest fragments with the other habitats of the landscape and represent a highly functional resource for the preservation of biodiversity that serves as a complement to but not a substitute for cloud forest in this notably modified landscape.     

The Potential for Species Conservation in Tropical Secondary Forests

Abstract: In the wake of widespread loss of old‐growth forests throughout the tropics, secondary forests will likely play a growing role in the conservation of forest biodiversity. We considered a complex hierarchy of factors that interact in space and time to determine the conservation potential of tropical secondary forests. Beyond the characteristics of local forest patches, spatial and temporal landscape dynamics influence the establishment, species composition, and persistence of secondary forests. Prospects for conservation of old‐growth species in secondary forests are maximized in regions where the ratio of secondary to old‐growth forest area is relatively low, older secondary forests have persisted, anthropogenic disturbance after abandonment is relatively low, seed‐dispersing fauna are present, and old‐growth forests are close to abandoned sites. The conservation value of a secondary forest is expected to increase over time, as species arriving from remaining old‐growth forest patches accumulate. Many studies are poorly replicated, which limits robust assessments of the number and abundance of old‐growth species present in secondary forests. Older secondary forests are not often studied and few long‐term studies are conducted in secondary forests. Available data indicate that both old‐growth and second‐growth forests are important to the persistence of forest species in tropical, human‐modified landscapes.     

Thermal ecology of the neotropical army ant Eciton burchellii.

I explored the thermal ecology of Eciton burchellii, a New World army ant, in primary forest and forest fragments in the Atlantic lowlands of Costa Rica in 2002 and 2003. My primary objective was to determine whether high surface temperatures in pastures surrounding forest fragments posed a thermal barrier to ant colonies within those fragments; secondarily, I assessed whether thermal gradients within continuous moist forest were sufficient to elicit avoidance reactions from foraging colonies. E. burchellii colonies in forest fragments avoided entering open pasture in full sun (51.3 degrees C) on 100% of all edge interactions; however, edges were readily crossed where artificial shaded areas had previously been installed. Ant raids in primary forest avoided artificially established temperatures >43 degrees C but tolerated 45.5 degrees C in the presence of prey baits. Captive ants held at 43 degrees C survived 18.5 min; at temperatures of 51.3 degrees C survival time was only 2.8 min. Ants running on established foraging trails increased running velocity by 18% when substrate temperature was raised from 28.4 degrees to 38.0 degrees C, and they abandoned trails at temperatures >43 degrees C. The standard deviation (s) of temperatures on active raid trails in continuous forest was 2.13 degrees C, while nearby systematic sampling revealed a greater background standard deviation of 4.13 degrees C. E. burchellii colonies in this region appear to be living surprisingly near their upper limits of thermal tolerance. The heat of open pastures alone is sufficient to prevent their exiting forest fragments, or entering similarly hot areas within continuous forest. Shaded vegetative corridors are sufficient to permit mobility between isolated fragments, and their preservation should be encouraged. Despite views that tropical lowland moist forests have an essentially homogenous microclimate, army ants appear to avoid local hot spots on the forest floor, steering daily foraging trails to follow cooler routes than would be expected by chance. As deforestation remakes tropical landscapes worldwide, it is important to consider the thermal consequences of these actions and their effect on keystone organisms such as army ants. Changes in global climate patterns are likely to affect even evergreen tropical systems whose organisms may be sensitive to finer microclimatic variation than previously suspected.     

Recovery of Faunal Communities During Tropical Forest Regeneration

Abstract:  As mature tropical forests are cleared, secondary forests may play an important role in the conservation of animal species, depending on how fast animal communities recover during forest regeneration. I reviewed published studies on the recovery of animal species richness and composition during tropical forest regeneration. In 38 of the 39 data sets I examined, conversion of forest to agriculture or pasture substantially reduced species richness. Given suitable conditions for forest recovery, the species richness of the animal taxa considered can be predicted to resemble that of mature forests roughly 20–40 years after land abandonment. At least for ants and birds, however, recovery of species composition appears to take substantially longer than recovery of species richness. Because species richness for many taxa appears to recover relatively rapidly in secondary forests, conservation of secondary forests may be an effective investment in future diversity. The slower recovery of species composition indicates, however, that some species will require stands of mature forest to persist.     

A Comparison of Logging Systems and Bat Diversity in the Neotropics

Abstract:  Evaluating logging systems to determine which are most compatible with the maintenance of biodiversity is of prime importance if tropical forests are to be managed in a sustainable way. Bats are model taxa for this purpose. Two different logging systems are used in the natural forest of the Victoria-Mayaro Forest Reserve in Trinidad: open range and periodic block. Open range is a continuous harvesting system and, in common with most methods used to log tropical forests worldwide, has few harvest controls other than girth limits for selected species. Periodic block is a polycyclic system, with felling based on ecological criteria assumed to be compatible with the maintenance of biodiversity. To compare the effects of periodic block and open-range systems on biodiversity, we determined bat species richness and abundance in each system and in primary forest. We caught bats in mist nets set at ground level and in the canopy and in harp traps. In total 1959 individuals representing 38 species were captured. Species richness was similar among primary forest and logged forest habitats, although bat diversity was lower in logged forest. The distributions of bat species abundance did not differ significantly between logged forest and primary forest. We found, however, that both logging systems lead to a decrease in gleaning animalivores and an increase in frugivores. The increase in frugivores was likely the result of an increase in the abundance of bat-dispersed pioneer fruiting plants in logged forest. Bats of periodic-block-managed forest were more similar to those of primary forest than those of forest logged using the open-range system, indicating that the periodic-block system is more compatible with the maintenance of bat diversity. Our results support the suggestion that the measured use of tropical forests can largely be compatible with biodiversity conservation.     

Effects of crowding due to habitat loss on species assemblage patterns

Terrestrial animals are negatively affected by habitat loss, which is assessed on a landscape scale, whereas secondary effects of habitat loss, such as crowding, are usually disregarded. Such impacts are inherently hard to address and poorly understood, and there is a growing concern that they could have dire consequences. We sampled birds throughout a deforestation process to assess crowding stress in an adjacent habitat remnant in the southern Brazilian Atlantic Forest. Crowding is expected of highly mobile taxa, especially given the microhabitat heterogeneity of Neotropical forests, and we hypothesized that the arrival of new individuals or species in refuges shifts assemblage patterns. We used point counts to obtain bird abundances in a before-after-control-impact design sampling of a deforestation event. Temporal changes in taxonomic and functional diversity were examined with metrics used to assess alpha and beta diversity, turnover of taxonomic and functional similarity, and taxonomic and functional composition. Over time increased abundance of some species altered the Simpson index and affected the abundance-distribution of traits in the habitat remnant. Taxonomic composition and functional composition changed in the remnant, and thus bird assemblages changed over time. Taxonomic and functional metrics indicated that fugitives affected resident assemblages in refuges, and effects endured >2 years after the deforestation processes had ceased. Dissimilarity of taxonomic composition between pre- and postdeforestation assemblages increased, whereas functional composition reverted to preimpact conditions. We found that ecological disruptions resulted from crowding and escalated into disruptions of species’ assemblages and potentially compromising ecosystem functioning. It is important to consider crowding effects of highly mobile taxa during impact assessments, especially in large-scale infrastructure projects that may affect larger areas than is assumed.     

Impact of cocoa agricultural intensification on bird diversity and community composition

To meet the growing demand for chocolate, cocoa (Theobroma cacao) agriculture is expanding and intensifying. Although this threatens tropical forests, cocoa sustainability initiatives largely overlook biodiversity conservation. To inform these initiatives, we analyzed how cocoa agriculture affects bird diversity at farm and landscape scales with a meta-analysis of 23 studies. We extracted 214 Hedges' g* comparisons of bird diversity and 14 comparisons of community similarity between a forest baseline and 4 farming systems that cover an intensification gradient in landscapes with high and low forest cover, and we summarized 119 correlations between cocoa farm features and bird diversity. Bird diversity declined sharply in low shade cocoa. Cocoa with >30% canopy cover from diverse trees retained bird diversity similar to nearby primary or mature secondary forest but held a different community of birds. Diversity of endemic species, frugivores, and insectivores (agriculture avoiders) declined, whereas diversity of habitat generalists, migrants, nectarivores, and granivores (agriculture associates) increased. As forest decreased on the landscape, the difference in bird community composition between forest and cocoa also decreased, indicating agriculture associates replaced agriculture avoiders in forest patches. Our results emphasize the need to conserve forested landscapes (land sparing) and invest in mixed-shade agroforestry (land sharing) because each strategy benefits a diverse and distinct biological community.     

Local and Landscape Correlates of Primate Distribution and Persistence in the Remnant Lowland Rainforests of the Upper Brahmaputra Valley,Northeastern India

Habitat fragmentation affects species distribution and abundance, and drives extinctions. Escalated tropical deforestation and fragmentation have confined many species populations to habitat remnants. How worthwhile is it to invest scarce resources in conserving habitat remnants within densely settled production landscapes? Are these fragments fated to lose species anyway? If not, do other ecological, anthropogenic, and species‐related factors mitigate the effect of fragmentation and offer conservation opportunities? We evaluated, using generalized linear models in an information‐theoretic framework, the effect of local‐ and landscape‐scale factors on the richness, abundance, distribution, and local extinction of 6 primate species in 42 lowland tropical rainforest fragments of the Upper Brahmaputra Valley, northeastern India. On average, the forest fragments lost at least one species in the last 30 years but retained half their original species complement. Species richness declined as proportion of habitat lost increased but was not significantly affected by fragment size and isolation. The occurrence of western hoolock gibbon (Hoolock hoolock) and capped langur (Trachypithecus pileatus) in fragments was inversely related to their isolation and loss of habitat, respectively. Fragment area determined stump‐tailed (Macaca arctoides) and northern pig‐tailed macaque occurrence (Macaca leonina). Assamese macaque (Macaca assamensis) distribution was affected negatively by illegal tree felling, and rhesus macaque (Macaca mulatta) abundance increased as habitat heterogeneity increased. Primate extinction in a fragment was primarily governed by the extent of divergence in its food tree species richness from that in contiguous forests. We suggest the conservation value of these fragments is high because collectively they retained the entire original species pool and individually retained half of it, even a century after fragmentation. Given the extensive habitat and species loss, however, these fragments urgently require protection and active ecological restoration to sustain this rich primate assemblage. Correlaciones Locales y de Paisaje de la Distribución y Persistencia de Primates en los Bosques Lluviosos Remanentes en el Valle del Alto Brahmaputra, Noreste de India     

Effects of Land‐Use Change on Community Composition of Tropical Amphibians and Reptiles in Sulawesi,Indonesia

Abstract: Little is known about the effects of anthropogenic land‐use change on the amphibians and reptiles of the biodiverse tropical forests of Southeast Asia. We studied a land‐use modification gradient stretching from primary forest, secondary forest, natural‐shade cacao agroforest, planted‐shade cacao agroforest to open areas in central Sulawesi, Indonesia. We determined species richness, abundance, turnover, and community composition in all habitat types and related these to environmental correlates, such as canopy heterogeneity and thickness of leaf litter. Amphibian species richness decreased systematically along the land‐use modification gradient, but reptile richness and abundance peaked in natural‐shade cacao agroforests. Species richness and abundance patterns across the disturbance gradient were best explained by canopy cover and leaf‐litter thickness in amphibians and by canopy heterogeneity and cover in reptiles. Amphibians were more severely affected by forest disturbance in Sulawesi than reptiles. Heterogeneous canopy cover and thick leaf litter should be maintained in cacao plantations to facilitate the conservation value for both groups. For long‐term and sustainable use of plantations, pruned shade trees should be permanently kept to allow rejuvenation of cacao and, thus, to prevent repeated forest encroachment.     

Two‐Stage Recovery of Amphibian Assemblages Following Selective Logging of Tropical Forests

There is a lack of quantitative information on the effectiveness of selective‐logging practices in ameliorating effects of logging on faunal communities. We conducted a large‐scale replicated field study in 3 selectively logged moist semideciduous forests in West Africa at varying times after timber extraction to assess post logging effects on amphibian assemblages. Specifically, we assessed whether the diversity, abundance, and assemblage composition of amphibians changed over time for forest‐dependent species and those tolerant of forest disturbance. In 2009, we sampled amphibians in 3 forests (total of 48 study plots, each 2 ha) in southwestern Ghana. In each forest, we established plots in undisturbed forest, recently logged forest, and forest logged 10 and 20 years previously. Logging intensity was constant across sites with 3 trees/ha removed. Recently logged forests supported substantially more species than unlogged forests. This was due to an influx of disturbance‐tolerant species after logging. Simultaneously Simpson's index decreased, with increased in dominance of a few species. As time since logging increased richness of disturbance‐tolerant species decreased until 10 years after logging when their composition was indistinguishable from unlogged forests. Simpson's index increased with time since logging and was indistinguishable from unlogged forest 20 years after logging. Forest specialists decreased after logging and recovered slowly. However, after 20 years amphibian assemblages had returned to a state indistinguishable from that of undisturbed forest in both abundance and composition. These results demonstrate that even with low‐intensity logging (≤3 trees/ha) a minimum 20‐year rotation of logging is required for effective conservation of amphibian assemblages in moist semideciduous forests. Furthermore, remnant patches of intact forests retained in the landscape and the presence of permanent brooks may aid in the effective recovery of amphibian assemblages. Recuperación de Ensambles de Anfibios en Dos Etapas Después de la Tala Selectiva de Bosques Tropicales     

Long-term patterns in tropical reforestation: plant community composition and aboveground biomass accumulation.

Primary tropical forests are renowned for their high biodiversity and carbon storage, and considerable research has documented both species and carbon losses with deforestation and agricultural land uses. Economic drivers are now leading to the abandonment of agricultural lands, and the area in secondary forests is increasing. We know little about how long it takes for these ecosystems to achieve the structural and compositional characteristics of primary forests. In this study, we examine changes in plant species composition and aboveground biomass during eight decades of tropical secondary succession in Puerto Rico, and compare these patterns with primary forests. Using a well-replicated chronosequence approach, we sampled primary forests and secondary forests established 10, 20, 30, 60, and 80 years ago on abandoned pastures. Tree species composition in all secondary forests was different from that of primary forests and could be divided into early (10-, 20-, and 30-year) vs. late (60- and 80-year) successional phases. The highest rates of aboveground biomass accumulation occurred in the first 20 years, with rates of C sequestration peaking at 6.7 +/- 0.5 Mg C x ha(-1) x yr(-1). Reforestation of pastures resulted in an accumulation of 125 Mg C/ha in aboveground standing live biomass over 80 years. The 80 year-old secondary forests had greater biomass than the primary forests, due to the replacement of woody species by palms in the primary forests. Our results show that these new ecosystems have different species composition, but similar species richness, and significant potential for carbon sequestration, compared to remnant primary forests.