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

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

A Flood Plain Palm Forest in the Luquillo Mountains of Puerto Rico Five Years After Hurricane Hugo1

Long-term studies are needed to understand the dynamics of tropical forests, particularly those subject to periodic disturbances such as hurricanes. We studied a flood plain Prestoea montana palm forest in the Luquillo Mountains of Puerto Rico over a 15-yr period (1980–1995), which included the passage of Hurricane Hugo in September 1989. The passage of the hurricane caused the dominant species to become more dominant and created low instantaneous tree mortality (1% of stems) and reductions in tree biomass (-16 Mg/ha/yr) and density, although not in basal area. Five years after the hurricane, the palm flood plain forest had exceeded its prehurricane aboveground tree biomass, tree density, and basal area. Aboveground tree biomass accumulated at a rate of 9.2 Mg/ha/yr, 76 percent of which was due to palms. Before the hurricane this rate was on the order of 3 Mg/ha/yr. Forest floor litter decreased to prehurricane levels (6.7 Mg/ha), within 5 yr, mostly due to the disappearance of woody litter. Thirteen tree species not represented in the canopy entered the forest by regeneration, and 2 species suffered almost 20 percent/yr mortality over a 5-yr period after the storm (floodplain average of 2%/yr). Delayed tree mortality was twice as high as instantaneous tree mortality after the storm and affected dicotyledonous trees more than it did palms. Regencration of dicotyledonous trees, palms, and tree ferns was influenced by a combination of factors including hydroperiod, light, and space. Redundancy Data Analysis showed that the area near the river channel was the most favorable for plant regeneration. Palm regeneration was higher in locations with longer hydroperiods, while regeneration of dicotyledonous trees was higher in areas with low risk of flooding. This study shows how a periodic disturbance provides long-term opportunities for species invasions and long-term ecosystem response at the patch scale of < 1 ha.     

Hypocrealean (Hypocreales,Ascomycota) Fungal Diversity in Different Stages of Tropical Forest Succession in Costa Rica1

The relationship between forest succession and microfungal diversity has been poorly studied. Fungi provide important ecosystem services that may deteriorate in deforested or highly disturbed forests. To determine the possible effects of deforestation and forest succession on microfungi, species diversity of hypocrealean fungi (Ascomycota) was compared in forest stands in Eastern Costa Rica representing three stages of succession: 1–2, 25–27 yr old, and an old growth forest. Species diversity in a second‐growth forest fragment surrounded by timber plantations and second‐growth forest was also compared to that of a stand surrounded by old growth forest. The results show that the overall diversity of hypocrealean fungi was inversely proportional to the age of the forest stand, and each family showed different successional trends. Clavicipitaceae was more diverse in the old‐growth forest and was positively related to the age of the forest stand. Nectriaceae was highly diverse in the 1‐ to 2‐yr‐old stand and less diverse in the old‐growth stand. Saprobic and plant pathogenic fungal species were more diverse in the 1‐ to 2‐yr‐old stand and their diversity was inversely proportional to the age of the forest stand. The diversity of insect pathogens was positively related to the age of the forest stand. The 20‐ to 22‐yr‐old forest fragment had the lowest number of species overall. Based on the data gathered in this study, hypocrealean fungal species diversity is related to the successional stage and fragmentation of tropical forest.     

Plant Community Structure in Tropical Rain Forest Fragments of the Western Ghats,India1

Changes in tree, liana, and understory plant diversity and community composition in five tropical rain forest fragments varying in area (18–2600 ha) and disturbance levels were studied on the Valparai plateau, Western Ghats. Systematic sampling using small quadrats (totaling 4 ha for trees and lianas, 0.16 ha for understory plants) enumerated 312 species in 103 families: 1968 trees (144 species), 2250 lianas (60 species), and 6123 understory plants (108 species). Tree species density, stem density, and basal area were higher in the three larger (> 100 ha) rain forest fragments but were negatively correlated with disturbance scores rather than area per se. Liana species density, stem density, and basal area were higher in moderately disturbed and lower in heavily disturbed fragments than in the three larger fragments. Understory species density was highest in the highly disturbed 18‐ha fragment, due to weedy invasive species occurring with rain forest plants. Nonmetric multidimensional scaling and Mantel tests revealed significant and similar patterns of floristic variation suggesting similar effects of disturbance on community compositional change for the three life‐forms. The five fragments encompassed substantial plant diversity in the regional landscape, harbored at least 70 endemic species (3.21% of the endemic flora of the Western Ghats–Sri Lanka biodiversity hotspot), and supported many endemic and threatened animals. The study indicates the significant conservation value of rain forest fragments in the Western Ghats, signals the need to protect them from further disturbances, and provides useful benchmarks for restoration and monitoring efforts.     

Influence of Forest Type and Tree Species on Canopy‐Dwelling Beetles in Budongo Forest,Uganda1

Beetles were collected on 64 trees of four species (Cynometra alexandri C. H. Wright, Rinorea beniemis (Welwitsch ex Olivier) Kuntze, Teclea nobilis Delile, and Trichilia rubescens Olivier) in Budongo Forest, Uganda, using an insecticidal fogging technique. Selected tree species were abundant, taxonomically not closely related, and different in the shape of leaves, growth form, and size, with heights between 7 and 35 m. Trees were fogged in an old primary forest stand, in an area of secondary forest where selective logging was performed, and in a swamp forest. Eight conspecific trees per forest type were fogged. A total of 29,736 beetles were collected from all trees that could be assigned to 1433 (morpho)‐species; 41.6 percent were singletons and 89.6 percent of species were found with less than ten individuals. Abundant beetle taxa included Latridiidae (N= 4093), Chrysomelidae (3952), Staphylinidae (2931), Apioninae (2621), and Curculionidae (2457). Most species‐rich groups were Staphylinidae (N= 196 spp.), Curculionidae (189), and Chrysomelidae (148). Abundance increased in the order: primary < secondary < swamp forest. Due to the relatively high dominance of some species in the secondary forest, species richness increased in the order: secondary < primary < swamp forest. Beta diversity measures and factor analysis showed distinct differences among forest types but higher similarity of beetle communities on different tree species within one forest type. The taxonomic distribution of beetles in the secondary forest was more heterogeneous than in the primary forest. Analyses of the data revealed low host specificity even for phytophagous beetles, underlining the importance of habitat structure and chance effects on the spatial distribution of beetles in the canopy of Budongo Forest.     

The effect of forest disturbance on the leaf litter ant fauna in Ghana

In order to observe the effect of forest loss on the leaf litter ant fauna in Ghana, West Africa, samples were taken in primary forest, secondary forest and in cocoa plots. Ants were extracted from the leaf litter by sieving followed by suspension in Winkler bags. The species composition and species richness in the three different habitats were compared and no significant difference was found between them. It was concluded that most primary forest leaf litter ant species continue to survive in parts of the agricultural landscape which has largely replaced their original habitat.     

Identifying forest‐obligate birds in tropical moist cloud forest of Andean Ecuador

ABSTRACT Large‐scale transformation of forested landscapes is a major factor in loss of biological diversity in the American tropics. Investigators examining the responses of species to deforestation rarely control for variation in the amount of forest relative to other habitats at the landscape‐level. Bellavista Reserve on the western slope of the Andes in Ecuador is located between similar‐sized areas of pristine, protected forest, and deforested landscapes. We used strip‐transect counts and mist netting to evaluate habitat use by passerine birds in a habitat mosaic consisting of abandoned pastures, forest edges, forest fragments, and large blocks of interior tropical montane cloud forest (TMCF). During 3600 net hours, we had 1476 captures, including 346 recaptures. Of 78 species captured in mist nets, 30 had sufficient counts for Poison Rate Regression (PRR) modeling (a statistical method for comparing counts). Twelve species (40%) had capture patterns indicative of an affinity for mature TMCF, and 6 species (20%) had significantly higher counts in degraded areas (forest edge, forest fragment, and regenerating pastures) than in interior TMCF. The remaining 40% showed no significant bias in detection among habitats. Combined with strip‐count data, our results suggest that about 38% of the 119 species sampled at the Bellavista Reserve occur primarily in mature TMCF, avoiding edges and early second‐growth forest. Populations of these species may be vulnerable to further loss, fragmentation, and degradation of TMCF and, as such, deserve additional study and a place on lists of species of conservation concern.     

Land‐use controls on carbon biogeochemistry in lowland streams of the Congo Basin

The flux and composition of carbon (C) from land to rivers represents a critical component of the global C cycle as well as a powerful integrator of landscape‐level processes. In the Congo Basin, an expansive network of streams and rivers transport and cycle terrigenous C sourced from the largest swathe of pristine tropical forest on Earth. Increasing rates of deforestation and conversion to agriculture in the Basin are altering the current regime of terrestrial‐to‐aquatic biogeochemical cycling of C. To investigate the role of deforestation on dissolved organic and inorganic C (DOC and DIC, respectively) biogeochemistry in the Congo Basin, six lowland streams that drain catchments of varying forest proportion (12%–77%) were sampled monthly for 1 year. Annual mean concentrations of DOC exhibited an asymptotic response to forest loss, while DIC concentrations increased continuously with forest loss. The isotopic signature of DIC became significantly more enriched with deforestation, indicating a shift in source and processes controlling DIC production. The composition of dissolved organic matter (DOM), as revealed by ultra‐high‐resolution mass spectrometry, indicated that deforested catchments export relatively more aliphatic and heteroatomic DOM sourced from microbial biomass in soils. The DOM compositional results imply that DOM from the deforested sites is more biolabile than DOM from the forest, consistent with the corresponding elevated stream CO₂ concentrations. In short, forest loss results in significant and comprehensive shifts in the C biogeochemistry of the associated streams. It is apparent that land‐use conversion has the potential to dramatically affect the C cycle in the Congo Basin by reducing the downstream flux of stable, vascular‐plant derived DOC while increasing the transfer of biolabile soil C to the atmosphere.     

Inner‐crown Microenvironments of Two Emergent Tree Species in a Lowland Wet Forest1

Vascular epiphyte communities, comprising up to 25 percent of tropical forest flora, contribute to plant diversity and thus ecosystem‐level processes; however, one of the proximal determinants of those communities, microclimate, is little studied. Here we present the first comprehensive study of microclimates in the inner crowns of two emergent tree species, Hyeronima alchorneoides and Lecythis ampla, at La Selva Biological Station, Costa Rica. We examined photon flux density, temperature, vapor pressure, and humidity in inner‐crown branches during the wet and dry seasons and during the wet‐season leafless phase of Lecythis. In both seasons, the percentage daily PFD in foliated Lecythis crowns (9%, wet season; 11%, dry season) was significantly higher than in Hyeronima crowns (5%, both seasons), with the leafless wet‐season PFD of Lecythis reaching 23 percent of full sun. Temperature and vapor pressure varied less in Hyeronima than in Lecythis crowns during the dry season. Microenvironmental conditions for epiphytes within Hyeronima crowns were more spatially and temporally homogeneous and were more buffered from ambient conditions than within Lecythis crowns. Growing conditions within the crowns of the same trees and among different trees were measurably different and are likely to affect the structure and composition of the resident epiphyte communities.     

Genetic Divergence During Long‐term Isolation in Highly Diverse Populations of Tropical Trees Across the Eastern Arc Mountains of Tanzania

The Eastern Arc Mountains are believed to support some of the oldest tropical forest in the world. The current distribution of this forest is highly fragmented due to a combination of long‐term effects of past changes in global climate and more recent deforestation. We sought to explore the hypothesized antiquity and long‐term isolation of the Eastern Arc montane forests based on an assessment of the geographical distribution and interspecies similarity of chloroplast DNA sequence variation in five forest trees. Data were used to investigate regional patterns of diversity and population structure based on intraspecific phylogenies, and results were interpreted against hypotheses on ecosystem age and connectivity. Regional diversity was high, with up to 22 chloroplast DNA haplotypes being recorded within a species across the sampled populations. Geographical concordance of genetic and geographic structure was weak to absent in all species and there was little similarity of genetic structure between species. Haplotype sharing between mountain blocks was extremely limited. The generally weak phylogeographical structure, in conjunction with high regional diversity and genetic uniqueness of individual mountain forests does not support the assumption of widespread genetic connectivity of the mountain forests, indicating instead a pattern of past isolation and ongoing diversification. Our findings substantially add to understanding patterns of diversity in this region and lend weight to calls to use more sophisticated biodiversity assessments when setting regional conservation and research funding priorities.     

Differences in the Effects of Selective Logging on Amphibian Assemblages in Three West African Forest Types

Making generalizations about the impact of commercial selective logging on biodiversity has so far remained elusive. Species responses to logging depend on a number of factors, many of which have not been studied in detail. These factors may include the natural forest conditions (forest types) under which logging impacts are investigated; but this question has so far remained unexamined. In a large‐scale replicate study we aimed at clarifying the relationship between logging and forest types on leaf litter frogs. We contrast three distinct and naturally occurring forest types, including wet evergreen, moist evergreen and semi‐deciduous forests. Selectively logged sites were compared with primary forest sites for each forest type. We found that the response of frog communities to logging varies in different forest types. In the wet evergreen forest, richness was higher in logged forest than primary forest, while diversity measures were not different between logged and primary forest habitats. In the moist evergreen, richness and diversity were higher in selectively logged areas compared with primary forest habitats. In the semi‐deciduous, logged forests were characterized by drastic loss of forest specialists, reduced richness, and diversity. These results indicate that the net effect of logging varies with respect to forest type. Forest types that are characterized by adverse climatic conditions (i.e., low rainfall and protracted dry seasons) are more likely to produce negative effects on leaf litter anuran communities. For comparisons of the impact of logging on species to be effective, future research must endeavor to include details of forest type.     

Land‐use strategies to balance livestock production,biodiversity conservation and carbon storage in Yucatán,Mexico

Balancing the production of food, particularly meat, with preserving biodiversity and maintaining ecosystem services is a major societal challenge. Research into the contrasting strategies of land sparing and land sharing has suggested that land sparing—combining high‐yield agriculture with the protection or restoration of natural habitats on nonfarmed land—will have lower environmental impacts than other strategies. Ecosystems with long histories of habitat disturbance, however, could be resilient to low‐yield agriculture and thus fare better under land sharing. Using a wider suite of species (birds, dung beetles and trees) and a wider range of livestock‐production systems than previous studies, we investigated the probable impacts of different land‐use strategies on biodiversity and aboveground carbon stocks in the Yucatán Peninsula, Mexico—a region with a long history of habitat disturbance. By modelling the production of multiple products from interdependent land uses, we found that land sparing would allow larger estimated populations of most species and larger carbon stocks to persist than would land sharing or any intermediate strategy. This result held across all agricultural production targets despite the history of disturbance and despite species richness in low‐ and medium‐yielding agriculture being not much lower than that in natural habitats. This highlights the importance, in evaluating the biodiversity impacts of land use, of measuring population densities of individual species, rather than simple species richness. The benefits of land sparing for both biodiversity and carbon storage suggest that safeguarding natural habitats for biodiversity protection and carbon storage alongside promoting areas of high‐yield cattle production would be desirable. However, delivering such landscapes will probably require the explicit linkage of livestock yield increases with habitat protection or restoration, as well as a deeper understanding of the long‐term sustainability of yields, and research into how other societal outcomes vary across land‐use strategies.     

Preferences and Food Quality of Fruit‐Feeding Butterflies in Kibale Forest,Uganda1

Many butterflies in tropical forests feed on fruits that have fallen to the forest floor. This substrate differs in many ways from floral nectar, and therefore fruit‐feeding butterflies are expected to possess adaptations for efficient foraging, choice, and ingestion of their food. Differences in food quality and in spatial and temporal availability are also likely to have led to life history evolution. Here we describe the sugar and nitrogen content of fruits that butterflies feed on in a tropical forest in Uganda, and measure the attractiveness of these fruits to the local butterfly fauna together with the role that decay plays in the attraction. These data are supplemented with feeding observations at fruit falls in the forest. Our results show that (1) fruits contain significant and variable concentrations of sugar and nitrogen, and constitute a nutritious food source for butterflies in tropical forests; (2) fruit‐feeding butterflies use cues from the fruits and fermentation products to locate their food; (3) different classes of fruit‐feeding butterflies may vary in their preferences for certain fruits, and differ in their ability to find preferred food; and (4) fruit choice is not strongly correlated with attractiveness or nutrient content. The results are discussed in the light of the evolution of food searching and life history strategies.     

Forest Succession in Tropical Hardwood Hammocks of the Florida Keys: Effects of Direct Mortality from Hurricane Andrew1

A tree species replacement sequence for dry broadleaved forests (tropical hardwood hammocks) in the upper Florida Keys was inferred from species abundances in stands abandoned from agriculture or other anthropogenic acitivities at different times in the past. Stands were sampled soon after Hurricane Andrew, with live and hurricane‐killed trees recorded separately; thus it was also possible to assess the immediate effect of Hurricane Andrew on stand successional status. We used weighted averaging regression to calculate successional age optima and tolerances for all species, based on the species composition of the pre‐hurricane stands. Then we used weighted averaging calibration to calculate and compare inferred successional ages for stands based on (1) the species composition of the pre‐hurricane stands and (2) the hurricane‐killed species assemblages. Species characteristic of the earliest stages of post‐agricultural stand development remains a significant component of the forest for many years, but are gradually replaced by taxa not present, even as seedlings, during the first few decades. This compositional sequence of a century or more is characterized by the replacement of deciduous by evergreen species, which is hypothesized to be driven by increasing moisture storage capacity in the young organic soils. Mortality from Hurricane Andrew was concentrated among early‐successional species, thus tending to amplify the long‐term trend in species composition.     

Long‐term Spatial Changes in the Distribution of the Brazilian Atlantic Forest

The aim of this study is to represent simultaneously changes in the spatial distribution of the Atlantic forest during the last 17,000 years. To characterize such changes, here we focused on three different forest physiognomies, evergreen, semi‐deciduous, and Araucaria, and we provide a list of indicator taxa for each class retrieved from the original published datasets. A review of the published fossil pollen records allowed us to classify regional behaviors in three main areas of distribution, north of 15°S, between 15° and 23°S and south of 23°S latitude that correspond to three climatic geographical barriers. Statistical probability density function method was used to illustrate changes in forest physiognomies throughout the three distribution areas. We show that the three modern barriers also functioned through the past. Asynchronous patterns of forest physiognomies are linked to an antiphasing pattern of monsoon precipitation between the northern and central area, whereas in the southern area, it is linked to the frequency and intensity of the polar advection in the subtropics. Our results attest to strong climate forcing on forest distribution between the late glacial and the interglacial period. They call into question the common reference to the last glacial maximum as a major (and sometimes as the only) driver of forest‐related vicariance and genetic diversity patterns, but suggest that instead, orbital cycles were the main drivers of the successive expansion/contraction of the Atlantic forest throughout the Quaternary.     

Vegetation Structure,Composition, and Species Richness Across a 56‐year Chronosequence of Dry Tropical Forest on Providencia Island,Colombia1

We compared vegetation structure and species richness across a 56‐yr chronosequence of six replicated age classes of dry tropical forest on the island of Providencia, Colombia, in the Southwest Caribbean. Stand age classes were determined using sequential, orthorectified panchromatic aerial photos acquired between 1944 and 1996 and Landsat 7 ETM + satellite imagery from 2000. Along the chronosequence we established 59 plots of 2 × 50 m (0.01 ha) to document changes in species richness, basal area, tree height, stem density, and sprouting ability. All woody trees and shrubs >2.5 cm diameter at breast height (DBH) were censused and measured. Although woody species density reached a peak in stands from 32 to 56 yr old, rarefaction analysis showed that species richness increased linearly with stand age and was highest in stands 56 yr old or greater. Nonparametric, abundance‐based estimators of species richness also showed positive and linear associations with age. Basal area and mean tree height were positively associated with age since abandonment, while sprouting ability showed a negative relationship. Our results indicate rapid recovery of woody species richness and structural characteristics along this tropical dry forest chronosequence.     

Incidence of Fungal Necrotrophic and Biotrophic Pathogens in Pioneer and Shade‐tolerant Tropical Rain Forest Trees

This study assessed the hypothesis that plant life history traits determine the incidence of fungal biotrophic and necrotrophic pathogens in pioneer vs. shade‐tolerant tropical plant species. Considering that pioneer species mainly invest in induced defenses, we expected a negative relationship between the incidence of biotrophic and necrotrophic pathogens; in contrast, as shade‐tolerant species invest heavily in constitutive defenses, we expected to find no correlation between the incidence of biotrophic and necrotrophic pathogens. These ideas were evaluated by assessing standing levels of fungal damage in a set of pioneer and shade‐tolerant species from the Lacandona tropical rain forest (Mexico). The results showed that among pioneer plant species, leaves with biotrophic lesions were between 34 and 44 percent more abundant than those with necrotic lesions. In contrast, among shade‐tolerant species, the proportions of leaves with necrotic lesions were 17–23 percent higher than those of leaves with injuries caused by biotrophic pathogens. Our study suggests that tropical tree species might present different defense strategies depending on the life‐style of the pathogens that attack them, and the life history strategy of the attacked host plant species. Thus, the host constitutive and induced defenses, as well as the mechanisms used by different types of pathogens to circumvent those defenses maybe responsible for the patterns of attack observed in perennial tropical plants. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .