Effects of Artificial Roosts for Frugivorous Bats on Seed Dispersal in a Neotropical Forest Pasture Mosaic

Abstract:  In the Neotropics ongoing deforestation is producing open and heavily fragmented landscapes dominated by agriculture, mostly plantations and cattle pastures. After some time agriculture often becomes uneconomical and land is abandoned. Subsequent habitat regeneration may be slow because seed inputs are restricted by a lack of incentives—such as suitable roost sites—for seed dispersers to enter deforested areas. Increasing environmental awareness has fostered growing efforts to promote reforestation. Practical and cost-efficient methods for kick-starting forest regeneration are, however, lacking. We investigated whether artificial bat roosts for frugivorous bat species can attract these key seed dispersers to deforested areas, thereby increasing seed rain. We installed artificial bat roosts in a forest-pasture mosaic in the Costa Rican Atlantic lowlands and monitored bat colonization and seed dispersal. Colonization occurred within a few weeks of installation, and 10 species of bats occupied the artificial roosts. Five species of frugivorous or nectarivorous bats colonized artificial roosts permanently in both primary habitat and in deforested areas, in numbers similar to those found in natural roosts. Seed input around artificial roosts increased significantly. Sixty-nine different seed types, mostly of early-successional plant species, were transported by bats to artificial roosts in disturbed habitats. The installation of artificial bat roosts thus successfully attracted frugivorous bats and increased seed inputs into degraded sites. This method is likely to speed up early-vegetation succession, which in turn will attract additional seed dispersers, such as birds, and provide a microhabitat for seeds of mid- and late-successional plants. As well as supporting natural forest regeneration and bat conservation, this cost-efficient method can also increase environmental awareness among landowners.     

Associations of Forest Cover,Fragment Area,and Connectivity with Neotropical Understory Bird Species Richness and Abundance

Theoretical and empirical studies demonstrate that the total amount of forest and the size and connectivity of fragments have nonlinear effects on species survival. We tested how habitat amount and configuration affect understory bird species richness and abundance. We used mist nets (almost 34,000 net hours) to sample birds in 53 Atlantic Forest fragments in southeastern Brazil. Fragments were distributed among 3 10,800‐ha landscapes. The remaining forest in these landscapes was below (10% forest cover), similar to (30%), and above (50%) the theoretical fragmentation threshold (approximately 30%) below which the effects of fragmentation should be intensified. Species‐richness estimates were significantly higher (F= 3715, p = 0.00) where 50% of the forest remained, which suggests a species occurrence threshold of 30–50% forest, which is higher than usually occurs (<30%). Relations between forest cover and species richness differed depending on species sensitivity to forest conversion and fragmentation. For less sensitive species, species richness decreased as forest cover increased, whereas for highly sensitive species the opposite occurred. For sensitive species, species richness and the amount of forest cover were positively related, particularly when forest cover was 30–50%. Fragment size and connectivity were related to species richness and abundance in all landscapes, not just below the 30% threshold. Where 10% of the forest remained, fragment size was more related to species richness and abundance than connectivity. However, the relation between connectivity and species richness and abundance was stronger where 30% of the landscape was forested. Where 50% of the landscape was forested, fragment size and connectivity were both related to species richness and abundance. Our results demonstrated a rapid loss of species at relatively high levels of forest cover (30–50%). Highly sensitive species were 3‐4 times more common above the 30–50% threshold than below it; however, our results do not support a unique fragmentation threshold. Asociaciones de la Cobertura Forestal, Superficie del Fragmento y Conectividad con la Riqueza y Abundancia de Aves Neotropicales de Sotobosque     

Effects of Forest Fragmentation on Seed Dispersal and Seedling Establishment in Ornithochorous Trees

Abstract: Habitat fragmentation increases seed dispersal limitation across the landscape and may also affect subsequent demographic stages such as seedling establishment. Thus, the development of adequate plans for forest restoration requires an understanding of mechanisms by which fragmentation hampers seed delivery to deforested areas and knowledge of how fragmentation affects the relationship between seed‐deposition patterns and seedling establishment. We evaluated the dispersal and recruitment of two bird‐dispersed, fleshy‐fruited tree species (Crataegus monogyna and Ilex aquifolium) in fragmented secondary forests of northern Spain. Forest fragmentation reduced the probability of seed deposition for both trees because of decreased availability of woody perches and fruit‐rich neighborhoods for seed dispersers, rather than because of reductions in tree cover by itself. The effects of fragmentation went beyond effects on the dispersal stage in Crataegus because seedling establishment was proportional to the quantities of bird‐dispersed seeds arriving at microsites. In contrast, postdispersal mortality in Ilex was so high that it obscured the seed‐to‐seedling transition. These results suggest that the effects of fragmentation are not necessarily consistent across stages of recruitment across species. Habitat management seeking to overcome barriers to forest recovery must include the preservation, and even the planting, of fleshy‐fruited trees in the unforested matrix as a measure to encourage frugivorous birds to enter into open and degraded areas. An integrative management strategy should also explicitly consider seed‐survival expectancies at microhabitats to preserve plant‐population dynamics and community structure in fragmented landscapes.     

Geographic Range Fragmentation and Abundance in Neotropical Migratory Birds

Populations of neotropical migrant landbirds have experienced significant declines in recent years. We investigated potential consequences of these declines by examining the relationship between abundance and fragmentation of geographic ranges of species on the North American breeding grounds. We estimated areograpbic fragmentation using the box dimension of a species' geographic range and demographic fragmentation using the fractal dimension of the semivariance function calculated from samples of population abundance across species' geographic ranges. We found a negative relationship between average abundance and demographic fragmentation for neotropical migrants, but not for residents. We also showed that demographic fragmentation and areographic fragmentation are inversely related for residents, but not for neotropical migrants. These results imply that neotropical migrants may be more sensitive to extinction than are residents.     

Rarity in Neotropical Forest Mammals Revisited

The identification of rare species is an important goal in conservation biology. Recent attempts to classify rare species have emphasized dichotomies in such characteristics as local population density, area of distribution, and degree of ecological specialization. In particular, Arita et al. (1990) dichotomized 100 Neotropical forest mammals according to local population density and area of distribution. Among these species of mammals, mean body mass was significantly associated with local population density and area of distribution. We argue that the effects of body mass should be removed before species are classified with respect to rarity. We re-evaluated the data on Neotropical mammal species, using regression analyses to remove the effects of body mass on population density and area of distribution, followed by analysis of residuals. This new method resulted in substantial changes in the dichotomous classification of rare species. We combined the analysis of regression residuals with a ranking procedure that assumed that local population density and area of distribution were equally important in their effects on rarity. The new ranking technique produced another different classification of the rarity of the Neotropical forest mammal species. A graphical analysis showed that ranked species differed substantially in their degree of rarity, and in the importance of local population density, area of distribution, or both, to their degree of rarity. The ranking method allows the species of greatest concern to be singled out, it can be modified to include additional variables such as niche breadth, and it should be helpful for making conservation decisions.     

Dispersal and social organization in the Neotropical Grey sac-winged bat Balantiopteryx plicata

Recent evidence suggests that tropical bats may frequently depart from the predominant mammalian male-biased dispersal pattern. So far, two emballonurid bat species that are closely related to our study species (Grey sac-winged bat, Balantiopteryx plicata) have been found to exhibit exceptional female-biased dispersal that is in accordance with father–daughter inbreeding avoidance. In contrast, using a combination of long-term behavioral observations of banded bats and DNA sequencing of the mitochondrial d-loop, our results suggest that B. plicata is the first Neotropical emballonurid with female philopatry and frequent male dispersal. However, just like in the other emballonurids, the age of females at first conception fell below the tenure of males. Thus, philopatric B. plicata females might face a father–daughter inbreeding risk if mating with males from their roosts. Such risk could be circumvented if mating occurs outside the nursing roost, e.g., in male mating aggregations. In contrast to other Neotropical emballonurid bats, the Grey sac-winged bat forms colonies with a greatly male-biased sex ratio (only 5 and 21 % females). Males of such colonies showed high roost fidelity and the sex ratio did not change throughout the year and for up to many years, suggesting an important role in mating. We conclude that studying the diverse mating and dispersal patterns of Neotropical emballonurids shows great potential to enlarge our understanding on how the proposed ultimate causes (i.e., avoidance of inbreeding and local competition between kin) affect the evolution of sex-biased dispersal.     

Bat Diversity and Abundance as Indicators of Disturbance in Neotropical Rainforests

Abstract: Evaluating the degree of disturbance of any region to determine its relative importance for conservation purposes requires procedures that are relatively inexpensive and that yield accurate results fast. Because bats are abundant, diverse, and easy to sample, especially in the Neotropical rainforest, they fulfill several of the requirements of indicator species as identified in the literature. For 10 months we sampled bat communities in the Selva Lacandona in Chiapas, Mexico, at 15 sites representing five habitats. We also measured 10 variables representing vegetation structure and diversity at each site. With fuzzy-set techniques we produced a gradient classification of disturbance for the 15 sites based on the vegetation data. We explored the relationship between vegetation conditions, described as the membership degrees in the construct "fuzzy forest set" (the complementary fuzzy set of "disturbance"), and four bat community variables. Bat species richness, number of rare bat species, and the bat diversity index were positively correlated with the vegetation scores, and relative abundance of the most abundant bat species was negatively correlated with vegetation scores. A high number of phyllostomine species in a community is a good indicator of low levels of disturbance. Although a single indicator group will probably not be sufficient for decision-making processes in conservation, evaluating bat populations may be a good first step in assessing an area's conservation value, especially in rainforest regions.     

Hunting Increases Dispersal Limitation in the Tree Carapa procera, a Nontimber Forest Product

Abstract:  The sustainability of seed extraction from natural populations has been questioned recently. Increased recruitment failure under intense seed harvesting suggests that seed extraction intensifies source limitation. Nevertheless, areas where more seeds are collected tend to also have more intense hunting of seed-dispersing animals. We studied whether such hunting, by limiting disperser activity, could cause quantitative dispersal limitation, especially for large crops and for crops in years of high seed abundance. In each of four Carapa procera (Meliaceae) populations in French Guiana and Surinam, two with hunting and two without, we compared seed fate for individual trees varying in crop size in years of high and low population-level seed abundance. Carapa seeds are a nontimber forest product and depend on dispersal by scatter-hoarding rodents for survival and seedling establishment. Hunting negatively affected the proportion of seeds dispersed and caused greater numbers of seeds to germinate or be infested by moths below parent trees, where they would likely die. Hunting of seed-dispersing animals disproportionally affected large seed crops, but we found no additional effect of population-level seed abundance on dispersal rates. Consistently lower rates of seed dispersal, especially for large seed crops, may translate to lower levels of seedling recruitment under hunting. Our results therefore suggest that the subsistence hunting that usually accompanies seed collection is at the cost of seed dispersal and may contribute to recruitment failure of these nontimber forest products. Seed extraction from natural populations may affect seedling recruitment less if accompanied by measures adequately incorporating and protecting seed dispersers.     

Disturbance-Mediated Mammal Persistence and Abundance-Area Relationships in Amazonian Forest Fragments

Abstract:  Few researchers have investigated the synergistic effects of tropical forest fragmentation and disturbance on species persistence and abundance. We examined effects of both forest-patch metrics and forest disturbance in determining richness and abundance of midsized to large-bodied mammal species in a highly fragmented Amazonian forest landscape. Twenty-one forest fragments, ranging from 2 to 14,480 ha, and two continuous forest sites were sampled based on sightings, tracks, line-transect censuses, armadillo burrow censuses, and camera trapping. Patch occupancy of 37 species recorded ranged from 4% to all forest sites surveyed. Forest fragment size was the strongest predictor of species persistence, explaining 90% of the variation in species richness. Information-theoretic analysis confirmed that fragment area was the most important explanatory variable for the overall species richness and abundance of mammal species, followed by surface fires, which affected the abundance of seven species. Large mammal species were typically absent from fragments <100 ha, whereas some ubiquitous species were favored by fragmentation, exhibiting hyperabundance in small patches. Our findings highlight the importance of large (>10,000 ha), relatively undisturbed forest patches to maximize persistence and maintain baseline abundances of Neotropical forest mammal species.     

Periodic Migration and Lowland Forest Refugia in a "Sedentary" Neotropical Bird, Wetmore's Bush-Tanager

Although the concept of tropical birds as sedentary is pervasive, evidence suggests many are not. Our grasp of movement status in tropical birds is decidedly poor, but the successful long-term conservation of these birds depends on such information. Sedentariness will likely doom much tropical avian diversity, but increased vagility is a two-edged sword: beneficial in promoting immigration, but detrimental in that more than one habitat may be required. Birds requiring more than one habitat may be unable to locate a particular type as landscape modifications increase. Our long-term data set from the Sierra de Los Tuxtlas in southern Veracruz, México, reveals infrequent, large-scale movements in a local highland endemic. Wetmore's Bush-Tanager ( Chlorospingus ophthalmicus wetmorei) seems occasionally dependent upon lowland forests (now greatly diminished) as a refugium from temporarily unsuitable highlands. Our data and observations lead us to three conclusions: 1) assumptions of sedentariness in tropical birds should be made with extreme caution; 2) normal, but periodic phenomena may be easily overlooked, even in relatively long-term studies; and 3) missing such phenomena jeopardizes the success of any conservation plan because over the long term a population may be dependent upon refugia only occasionally occupied.     

Forest Fragmentation Increases Nest Predation in the Eurasian Treecreeper

Abstract:  We used long-term breeding data to monitor the influences of fragmentation and habitat composition at different spatial scales on the reproductive success of Eurasian Treecreepers ( Certhia familiaris ) breeding in nest boxes. We collected data from the same forest patches (2.7–65.1 ha in size) during seven breeding seasons. Nest predation varied considerably over the years and was the primary cause of nesting failure (mean annual rate of 21.6 ± 12.8%). Nest predation explained most of the variation in fledgling production during the study period. Landscape-level fragmentation (radius of 500 m from territory center) affected nest predation more than did fragmentation on the territory scale (radius of 200 m from territory center). In general, nest loss due to predation in fragmented landscapes (32.4%) was almost threefold that of less fragmented (12.0%) landscapes. Of the habitat variables, predation rate correlated positively with the density of edges between forest and open land and with the proportion of sapling stands on the spatial scale of 500 m around a nest. In the core area of a territory (radius of 30 m from territory center), a high density of trees increased the frequency of nest predation. Further, a high proportion of agricultural land close to a nest site increased nest losses of treecreepers, probably because of a high degree of mustelid predation. Our results showed that the spatial scale on which we examined nest predation is important and that even within moderately fragmented landscapes it is possible to detect fragmentation-related nest predation.     

Seed predation by Neotropical rain forest mammals increases diversity in seedling recruitment

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.