Lemur responses to edge effects in the Vohibola III classified forest, Madagascar

Forest edges are dynamic zones characterized by the penetration (to varying depths and intensities) of conditions from the surrounding environment (matrix) into the forest interior. Although edge effects influence many tropical organisms, they have not been studied directly in primates. Edge effects are particularly relevant to lemurs because of the highly fragmented forest landscapes found in Madagascar. In this study, data are presented regarding how the densities of six lemur species (Avahi laniger, Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, Microcebus rufus, and Propithecus diadema edwardsi) varied between six 500-m interior transects and six 500-m edge transects in the Vohibola III Classified Forest in SE Madagascar. Diurnal (n = 433) and nocturnal (n = 128) lemur surveys were conducted during June-October 2003 and May-November 2004. A. laniger, E. rubriventer, and H. g. griseus exhibited a neutral edge response (no differences in densities between habitats). M. rufus and P. d. edwardsi had a positive edge response (higher densities in edge habitats), which may be related to edge-related variations in food abundance and quality. Positive edge responses by M. rufus and P. d. edwardsi may ultimately be detrimental due to edge-related anthropogenic factors (e.g., hunting by local people). The negative edge response exhibited by C. major (lower densities in edge habitats) may result from heightened ambient temperatures that inhibit torpor in edge habitats.     

片断化森林的边缘效应与自然保护区的设计管理

森林片断化引起了森林边缘物理环境与生物的变化。从森林边缘向林内,光辐射,温度、湿度、风等因素发生改变,这些反过 边缘上和边缘附近的生物产生极其重要的影响。有些研究表明,生物与非生物存在明显的边缘－核心的变化梯度,而有些则没有,除了影响边缘效应的客观因素（如边缘的取向、地势、年龄等）外,主观因素对这缘效应也有影响。边缘对生物与非生物的影响因片断化森林的面积、形状以及与其他森林片连接程度的不同而不同,     

High density of white-faced capuchins (Cebus capucinus) and habitat quality in the Taboga Forest of Costa Rica

Across the globe, primates are threatened by human activities. This is especially true for species found in tropical dry forests, which remain largely unprotected. Our ability to predict primate abundance in the face of human activity depends on different species' sensitivities as well as on the characteristics of the forest itself. We studied plant and primate distribution and abundance in the Taboga Forest, a 516-ha tropical dry forest surrounded by agricultural fields in northwestern Costa Rica. We found that the density of white-faced capuchins (Cebus capucinus) at Taboga is 2–6 times higher than reported for other long-term white-faced capuchin sites. Using plant transects, we also found relatively high species richness, diversity, and equitability compared with other tropical dry forests. Edge transects (i.e., within 100 m from the forest boundary) differed from interior transects in two ways: (a) tree species associated with dry forest succession were well-established in the edge and (b) canopy cover in the edge was maintained year-round, while the interior forest was deciduous. Sighting rates for capuchins were higher near water sources but did not vary between the edge and interior forest. For comparison, we also found the same to be true for the only other primate in the Taboga Forest, mantled howler monkeys (Alouatta palliata). Year-round access to water might explain why some primate species can flourish even alongside anthropogenic disturbance. Forest fragments like Taboga may support high densities of some species because they provide a mosaic of habitats and key resources that buffer adverse ecological conditions.     

Some observations on the present distribution and conservation of Alouatta and Ateles in Southern Mexico

Some information on the past and present distribution of Alouatta and Ateles in southern Mexico and on the impact of habitat destruction on their future existence is presented. Three surveys of primate populations and forest habitats were carried out between 1981 and 1983 in southeastern Mexico. Rapid destruction of the forest is reducing the number of forested areas where Alouatta and Ateles can exist and co-occur. Illegal hunting of the two primates also contributes to their rapid decline. Alouatta and Ateles live a precarious existence in a few large and small islands of forest, and efforts are needed to set aside some of these areas as reserves so that their future survival is ensured.     

Arthropods on plants in a fragmented Neotropical dry forest: A functional analysis of area loss and edge effects

Loss and fragmentation of natural ecosystems are widely recognized as the most important threats to biodiversity conservation, with Neotropical dry forests among the most endangered ecosystems. Area and edge effects are major factors in fragmented landscapes. Here, we examine area and edge effects and their interaction, on ensembles of arthropods associated to native vegetation in a fragmented Chaco Serrano forest. We analyzed family richness and community composition of herbivores, predators, and parasitoids on three native plant species in 12 fragments of varying size and at edge/interior positions. We also looked for indicator families by using Indicator Species Analysis. Loss of family richness with the reduction of forest fragment area was observed for the three functional groups, with similar magnitude. Herbivores were richer at the edges without interaction between edge and area effects, whereas predators were not affected by edge/interior position and parasitoid richness showed an interaction between area and position, with a steeper area slope at the edges. Family composition of herbivore, predator, and parasitoid assemblages was also affected by forest area and/or edge/interior situation. We found three indicator families for large remnants and five for edges. Our results support the key role of forest area for conservation of arthropods taxonomic and functional diversity in a highly threatened region, and emphasize the need to understand the interactions between area and edge effects on such diversity.     

Area and edge effects on leaf-litter decomposition in a fragmented subtropical dry forest

South American subtropical dry forests are highly threatened by fragmentation. Despite considerable research efforts aimed at predicting ecosystem alterations due to this driver of global change, we still need to deal with general principles to improve our ability to predict the impact of fragmentation. Our work is one of the few studies that analyse the relationship between forest fragmentation and decomposition. In 12 remnants of Chaco Serrano forest in Central Argentina we tested if decomposition rates of a common leaf-litter substrate varied with fragment size and between the forest edge and interior. Decomposition declined with fragment size, with no significant effects of location (edge/interior) or interaction between the two components of fragmentation. Our results suggest that in situ conditions for decomposition may change as a consequence of forest fragmentation, specifically as a result fragment size. This may lead to impaired nutrient recycling in smaller forest remnants.     

The capuchin, the howler, and the Caatinga: seed dispersal by monkeys in a threatened Brazilian forest

Regeneration of the Brazilian Caatinga forest may be restricted by the naturally low diversity and density of fruit-eating animals, which has been aggravated by local faunal extinction induced by human activities. We made a preliminary evaluation of the potential seed-dispersal role of capuchin (Cebus apella libidinosus) and howler monkeys (Alouatta caraya) in Serra da Capivara National Park. The monkeys dispersed at least 26 species. Alouatta dispersed larger seeds than Cebus, and the two species apparently dispersed seeds in different local habitats. Seed dispersal by monkeys potentially makes a significant contribution to Caatinga regeneration.     

Howling on the edge: Mantled howler monkey (Alouatta palliata) howling behaviour and anthropogenic edge effects in a fragmented tropical rainforest in Costa Rica

The function of long calling is a subject of interest across animal behaviour study, particularly within primatology. Many primate species have male‐specific long‐distance calls, including platyrrhines like the folivorous howler monkey (Alouatta spp.). Howler monkeys may howl to defend resources such as feeding trees or areas of rich vegetation from other monkey groups. This study tests the ecological resource defence hypothesis for howling behaviour in the mantled howler monkey (Alouatta palliata) and investigates how anthropogenic forest fragmentation may influence howling behaviour. More specifically, this study examines how howling bout rate, duration, precursors and tree species richness, DBH, and canopy cover vary in 100 m anthropogenic edge and interior forest zones at La Suerte Biological Research Station (LSBRS), a fragmented tropical rainforest in Costa Rica. Results show that tree species richness and canopy cover are higher in forest interior at this site, suggesting that monkeys should howl at greater rates in the interior to defend access to these higher‐quality vegetation resources. Overall, our results supported the ecological resource defence hypothesis. The main howl precursor was howling from neighbouring groups. Although howling rate did not differ between forest zones, howling bouts from forest interior were longer, had a greater number of howls per bout and were preceded by different precursors than howls from anthropogenic edge zones, including more howls from neighbouring groups. Our findings provide some of the first evidence for behavioural edge effects in primate vocal communication behaviour.     

Recovery of floristic diversity and basal area in natural forest regeneration and planted plots in a Costa Rican wet forest

When compared to planted reforestation, natural unassisted regeneration is often reported to result in slow recovery of biomass and biodiversity, especially early in succession. In some cases, naturally regenerating forests are not comparable to the community structure of primary forests after many decades. However, direct comparison of the outcomes of tropical forest restoration and natural regeneration is hindered by differences in metrics of forest recovery, inconsistency in land use histories, and dissimilarities in experimental design. We present the results of a replicated reforestation experiment comparing natural regeneration and polyculture tree planting at multiple diversity levels (3, 6, 9, or 12 native tree species), with uniform land use history and initial edaphic conditions. We compare the recovery of basal area and floristic diversity in these treatments after 5 yr of succession. Total basal area was higher in planted plots than in naturally regenerating plots, but it but did not vary among the different planted diversity levels. The basal area of woody recruits did not differ among treatments. The diversity of woody recruits increased substantially over time but did not vary among planting treatments. Species composition trajectories showed directional turnover over time, with no consistent differences among treatments. The convergence of restoration trajectories and similarity of floristic community diversity and composition across all treatments, after only 5 yr, provides evidence of the viability of natural regeneration for rapid restoration of forest biodiversity.     

Edge and shape effects on ant (Hymenoptera: Formicidae) species richness and composition in forest fragments

In this paper, we tested four hypotheses relative to edge and shape effects on ant communities: (i) forest edges have lower species richness than the remnant core; (ii) species richness increases with distance from the edge; (iii) irregularly shaped remnants have lower species richness than more regular remnants; (iv) there is a higher similarity of species composition between edge and core in irregular than in regular remnants. We sampled litter ant communities on the edge and core of ten remnants, in Viçosa, Minas Gerais, Brazil. Species richness was larger at the forest core than at the edges, although did not increase with distance from the edge. Species richness did not vary with shape complexity. The similarity of species composition between edge and core showed a decreasing trend with remnant area, and did not vary with shape complexity. The observed differences of species richness between forest core and edge may be due to higher harshness of edges, caused by environmental changes. The absence of relationship between species richness and distance from the edge might indicate the range of edge effects, which would be smaller than the smallest distance of core sampled. Therefore, edges would affect litter-dwelling ant species richness in a distance smaller than 50 m. The observation of species composition allowed us to notice an effect of fragmentation that would not be noticed if we were considering only species richness. Edge may serve as step to generalist species, which may use it to colonise forest remnants. Furthermore, small remnants are more colonisation-prone by such species, and have a more homogeneous species composition than large remnants.     

Small to modest impact of social group on the gut microbiome of wild Costa Rican capuchins in a seasonal forest

The horizontal transmission of pathogenic and beneficial microbes has implications for health and development of socially living animals. Social group is repeatedly implicated as an important predictor of gut microbiome structure among primates, with individuals in neighboring social groups exhibiting distinct microbiomes. Here we examine whether group membership is a predictor of gut microbiome structure and diversity across three groups of white‐faced capuchins (Cebus capucinus imitator) inhabiting a seasonal Costa Rican forest. We collected 62 fecal samples from 18 adult females during four sampling bouts. Sampling bouts spanned the dry‐to‐wet‐to‐dry seasonal transitions. To investigate gut microbial composition, we sequenced the V4 region of the 16S rRNA gene. We used the DADA2 pipeline to assign amplicon sequence variants and the RDP database to classify taxa. Our findings are: 1) gut microbiomes of capuchins clustered by social group in the late dry season, but this pattern was less evident in other sampling bouts; 2) social group was a significant variable in a PERMANOVA test of beta diversity, but it accounted for less variation than season; 3) social group was not an important predictor of abundance for the ten most abundant microbial taxa in capuchins; 4) when examining log2‐fold abundances of microbes between social groups, there were significant differences in some pairwise comparisons. While this is suggestive of group‐wide differences, individual variation may have a strong impact and should be assessed in future studies. Overall, we found a minor impact of social group membership on the gut microbiota of wild white‐faced capuchins. Future research including home range overlap and resource use, as well as fine‐scale investigation of individual variation, will further elucidate patterns of socially structured microbes.     

Understorey plant species richness and composition in metropolitan forest archipelagos: effects of forest size, adjacent land use and distance to the edge

Aim To address the relative role of adjacent land use, distance to forest edge, forest size and their interactions on understorey plant species richness and composition in perimetropolitan forests. Location The metropolitan area of Barcelona, north‐eastern Spain. Methods Twenty sampling sites were distributed in two forest size‐categories: small forest patches (8–90 ha) and large forest areas (> 18,000 ha). For each forest‐size category, five sites were placed adjacent to crops and five sites adjacent to urban areas. Vascular plant species were recorded and human frequentation was scored visually in 210 10 × 10 m plots placed at 10, 50 and 100 m from the forest edge, and additionally at 500 m in large forest areas. Plant species were grouped according to their ecology and rarity categories. A nonmetric multidimensional scaling (NMS) ordination was carried out to detect patterns of variation in species assemblage, and to explore the relationships between these patterns and the richness of the species groups and the studied factors. Factorial anovas were used to test the significance of the studied factors on the richness of species groups. Relationships between human frequentation and the studied variables were assessed through contingency tables. Results Forest‐size category was the main factor affecting synanthropic species (i.e. those thriving in man‐made or man‐disturbed habitats). Synanthropic species richness decreased with increasing distance from the forest edge and, when forests were adjacent to crops, it was higher in small forest patches than in large forest areas. Richness of rare forest species was lower in small forest patches than in large forest areas when forests were adjacent to urban areas. Richness of common forest species and of all forest species together were higher close to the forest edge than far from it when forests were adjacent to urban areas. Forests adjacent to urban areas were more likely to experience high human frequentation, particularly in those plots nearest to the forest edge. Main conclusions Forest‐size category and adjacent land use were the most important factors determining species richness and composition. The preservation of large forests adjacent to crops in peri‐urban areas is recommended, because they are less frequented by humans, are better buffered against the percolation of nonforest species and could favour the persistence of rare forest species.     

Substrate conditions,foliar nutrients and the distributions of two canopy tree species in a Costa Rican secondary rain forest

A 28 yr old secondary lowland rain forest in Costa Rica was dominated by two tree species of contrasting ecologies, Vochysia ferruginea - a species typical of well-drained but infertile soils of high Al saturation, and Cordia alliodora, a species requiring soils of at least moderate fertility. The two species exhibited markedly different spatial distributions in the forest and we sought to determine whether or not these different distributions were related to variation in substrate conditions and the nutrient requirements of the two species. Two soil types were present: Ultisols (Typic Haplohumults) and Inceptisols (Typic Dystropepts) and topography was of low hills. The study formed part of an analysis of site quality for timber production in the two species and therefore used standard forestry methods for such analyses. 36 plots of 20 m×20 m were established using predetermined criteria of site uniformity, presence of at least four dominant trees of at least one of the study species, and absence of disturbance. The abundance (number of individuals 10 cm dbh) of each species was determined in each plot. Soil samples were taken in 27 of the 36 plots, 9 plots being considered to replicate conditions already sampled, and 14 soil chemical and physical variables were measured using standard methods. Foliar nutrient analyses were carried out for trees of both species with crowns fully exposed to sunlight, distributed across the gradient of soil conditions. Variations in species abundances were not related to soil type, but plots in which Vochysia ferruginea was more abundant tended to be on steeper slopes with soils of higher exchangeable acidity and lower concentrations of Mn; Cordia alliodora was more abundant on gentler topography where soils had much lower exchangeable acidity but higher Mn. Differences between the two species in foliar nutrient concentrations were marked and supported previous interpretations of their nutrient requirements; foliar nutrients of Vochysia ferruginea were typical of tree species of moist tropical forest on infertile soils, and those of Cordia alliodora typical of a species requiring more fertile soils. Factors such as the distributions of seed trees at site abandonment may affect the spatial distributions of tree species in secondary forests such as that studied. The relationships of the distributions of the two species to substrate variation, however, and their evidently different nutrient requirements, support the hypothesis that variation in the composition and structure of the forest studied is at least partially related to exchangeable acidity and its dominant cation, Al.     

Fragment shape and tree species composition in tropical forests: a landscape level investigation

Fragmentation of tropical forest alters community composition as a result of changes in forest shape. This paper uses 22 hypotheses to test the effect of fragment shape on tree species composition in Ghana, West Africa, within biological categories of regeneration guild, rarity, phenology and dispersal. For both regenerating and mature trees, relationships between species composition and the shape of forest fragments were complex; almost half were significant but many failed to support the established hypotheses. Irregular shaped fragments had high proportions of regenerating, light‐demanding pioneers and mature, animal‐dispersed species. Species common to Ghana formed the foundation of communities in fragments of all shapes. Investigation at the landscape level indicated broad patterns of species change. Rigorous hypothesis testing is needed, following extensive demographic work on the ground, before population dynamics within tropical forest fragments can be comprehended fully and applied to conservation management.     

Influence of Edge Exposure on Tree Seedling Species Recruitment in Tropical Rain Forest Fragments1

Edge creation has a pronounced influence on the understory vegetation, but the effects of edges on seedling species recruitment are still poorly understood. In Central Amazonia, 9–19 years after fragmentation, we recorded species richness and net seedling recruitment rate in 1 ha blocks exposed to none, one, or multiple edges within forest fragments. One‐hectare blocks were located in the center (no edge), the edge (one edge), the corners (two edges) of 10 and 100 ha fragments, and in a 1 ha fragment (four edges). In 1991, we counted all tree seedlings 5–100 cm tall found within permanent 1 m² plots located within the 1 ha blocks. In May 1993, we manually removed all seedlings that were smaller than 1 m tall from the permanent plots. Six years and five months later (October 1999), all new seedlings recruited into the plots were counted and classified into distinct morphospecies. Species richness of recruited seedlings, scaled by total seedling density, declined from the center to the edge, the corner blocks, and then to the 1 ha fragment. Overall, the four‐edged, 1 ha fragment had the poorest species richness and the non‐edged 100 ha central block the highest. The total number of recruited individuals was 40 percent less than that previously present, with the 100 ha corner having the lowest recruitment. Pairwise comparisons showed that species similarity was related to edge number for the 100 and 1 ha fragments. Species rank/abundance curves showed that a subset of species was common in all blocks within the fragments, and that the 100 ha center held more rare species than any other 1 ha block. This study demonstrated that, in a given fragment patch, the number of tree seedling species recruited varied inversely with the number of edges.     

Natural vegetation cover in the landscape and edge effects: differential responses of insect orders in a fragmented forest

Human activities have led to global simplification of ecosystems, among which Neotropical dry forests are some of the most threatened. Habitat loss as well as edge effects may affect insect communities. Here, we analyzed insects sampled with pan traps in 9 landscapes (at 5 scales, in 100–500 m diameter circles) comprising cultivated fields and Chaco Serrano forests, at overall community and taxonomic order level. In total 7043 specimens and 456 species of hexapods were captured, with abundance and richness being directly related to forest cover at 500 m and higher at edges in comparison with forest interior. Community composition also varied with forest cover and edge/interior location. Different responses were detected among the 8 dominant orders. Collembola, Hemiptera, and Orthoptera richness and/or abundance were positively related to forest cover at the larger scale, while Thysanoptera abundance increased with forest cover only at the edge. Hymenoptera abundance and richness were negatively related to forest cover at 100 m. Coleoptera, Diptera, and Hymenoptera were more diverse and abundant at the forest edge. The generally negative influence of forest loss on insect communities could have functional consequences for both natural and cultivated systems, and highlights the relevance of forest conservation. Higher diversity at the edges could result from the simultaneous presence of forest and matrix species, although “resource mapping” might be involved for orders that were richer and more abundant at edges. Adjacent crops could benefit from forest proximity since natural enemies and pollinators are well represented in the orders showing positive edge effects.     

Chaco forest fragmentation effects on leaf litter decomposition are not explained by changes in litter fauna

Forest fragmentation is a component of global change, with substantial impact on biodiversity and ecosystem functioning. Despite extensive evidence of forest fragmentation effects on above‐ground ecological processes, little is understood about its below‐ground effects. Abundance and richness of leaf litter fauna can be affected by forest fragmentation, and this can have cascading effects on the decomposition process. Here, we examine how fragmentation of a subtropical dry forest affects aspects of ecosystem structure and functioning, by unravel area and edge effects on leaf litter fauna and decomposition rates and testing whether changes in abundance or richness of litter fauna mediated fragment area and edge effects on litter decomposition. We incubated litterbags filled with a common substrate, at the edge and interior of 12 fragments of Chaco Serrano forest in Central Argentina, for 180 days. We found that invertebrate abundance was higher at the forest edge but independent of fragment area, whereas decomposition declined with fragment size independently of edge or interior location. According to our results, the effect of forest size on decomposition was not mediated by changes in abundance or richness of leaf litter fauna, suggesting independent changes in ecosystem structure and functioning.     

Population of the black howler monkey (Alouatta pigra) in a fragmented landscape in Palenque,Chiapas, Mexico

Little is known about the population characteristics of Alouatta pigra under conditions of forest fragmentation-information that is important to understanding its tolerance to habitat loss. In this work we present data on forest loss and on troop size, age, and sex composition for a population of black howler monkeys existing in the fragmented landscape surrounding the Mayan site of Palenque, Chiapas, Mexico. Two aerial photos (1:70,000) of the study area (261 km(2)) taken in 1984 and 2001 were examined to assess forest loss. Between June and December 2001 and January and March 2002 we surveyed 44 forest fragments for the presence of howler monkeys. Examination of aerial photos showed that 33% of the forest present in 1984 had disappeared by 2001, and detected an increment in the number of forest fragments present in the landscape. We discovered a total of 115 howler monkeys living in 22 of the 44 forest fragments studied, of which 107 were members of 18 troops. The rest were solitary males or small groups of males living in isolated forest fragments. Troop size ranged from two to 15 individuals (mean 5.9+3.0 ind). 31% and 15% of individuals in the troops were juveniles and infants, respectively, suggesting continued reproductive activity. Howler monkey troops in the forest fragments were on average smaller (5.9+/-3.0 ind) than troops in the nearby protected forest of the Mayan site (7.0+/-2.8 ind). The mean density of howlers in the forest fragments was 119+/-82.9 ind/km(2). The establishment of corridors is suggested as a possible conservation scenario for the fragmented howler population investigated, and as a conservation measure to connect this population with the howler population found in the protected forest of the Mayan site.